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DECEMBER 30, 2005 • VOLUME 56 • NUMBERS 27-37 & INDEX 



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PROCEEDINGS 

OF THE 

CALIFORNIA ACADEMY OF SCIENCES 

(Fourth Series) 




CALIFORNIA 



ACADEMY OF SCIENCES 



San Francisco, California 



Copyright © 2005 by the California Academy of Sciences 

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Scientific Publications 

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Gary C. Williams, Ph.D., Associate Editor 

Michele L. Aldrich, Ph.D., Consulting Editor 

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From 1853 to 1906, when the Academy suffered the grievious loss of its handsome museum buildings as a result of 
the devastating earthquake and subsequent fire that swept San Francisco's downtown area, the Academy had been served 
by a number of exceptional persons as President of the fledgling organization. Each had distinguished himself as a scien- 
tist, mathematician, engineer, or in business, and most held administrative positions outside of the Academy. (Top row, left 
to right) Andrew Randall (1853-1856) had served as an assistant to Federal geologist David Dale Owen in the survey of 
the Wisconsin-Minnesota territories before coming to California, to Monterey; he was elected to the Califomia State 
Legislature. Leander Ransom (missing portrait; 1856-1866) after a successftil career as surveyor in Toledo, Ohio, came to 
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Surveyor General for Califomia. Ransom wisely selected the summit of Mount Diablo as the initial point for a survey and 
then, during the hot summer of 1851, he laid out the meridian and baseline markers needed to initiate the triangulation sur- 
vey of all of central Califomia and westem Nevada. Josiah Dwight Whitney (1867-1868) served for nearly a decade as head 
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entrepreneur, he served a terni as mayor of San Francisco, ser\'ed on the Board of Directors of the Bank of Califomia, and 
on numerous city commissions. Eusebius Molera (1905-1908), San Francisco engineer with an abiding interst in Aztec his- 
tory, also served as President Pro tern of the Academy's Board of Trustees. 

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 

Volume 56. No. 27. pp. 395-525 / '" \ December 30, 2005 




A Checklist and Preliminary BiBlrogrs^^y-tn the Recent, 

Freshwater Diatoms of Inland Environments 

of the Continental United States 



J.P. Kocioiek 

Diatom Collection, California Academy of Sciences 
875 Howard Street. San Francisco, CA 94103; Email: pkociolek® calacademy.org 

nJus puBUcation is dedicated to (Dr. 'Eugene jF. Stoermer an the occasion offus retirement. 

A checklist of freshwater diatoms of inland environments of the continental 
United States is compiled. It includes over 4500 names across almost 170 genera. The 
genus Navicula has the greatest number of taxa recorded, followed by Pinnularia, 
Nitzschia, Gomphonema, and Eunotia. A preliminary bibliography of reports of 
diatoms from U.S. inland environments is also presented. The number of biblio- 
graphic entries is over 1,200, and includes reports from diverse publications. It is 
hoped that the checklist and bibliography can be used to develop a comprehensive 
diatom flora of the United States. 



A challenge to documenting the Recent freshwater diatom flora of the United States is the lack 
of a synthesis of basic information. An understanding of the U.S. flora is important not only to 
assess the current state of our natural resources (in terms of biodiversity and subsequent changes 
due to changes in the physical, chemical and/or biological environment) but also to better under- 
stand the patterns of distribution within and outside the U.S. 

Kocioiek (submitted) has developed a list of taxa described from U.S. inland waters, and a 
comprehensive evaluation of all diatom names at the taxonomic level of species and below is forth- 
coming (Fourtanier & Kocioiek, based on the approach for genus names developed by Fourtanier 
& Kocioiek 1999, 2003). In addition to knowing what species have been described from the U.S., 
and the nomenclatural status of names, a third leg of the stool to begin a large-scale flora is a basic 
checklist of previously reported taxa. While many local and regional floras have been published 
fsee Kocioiek & Spaulding 2003 for a review of many of these), no comprehensive list of diatoms 
from the U.S. has been attempted in over 70 years (Boyer 1927a, b). 

To develop a checklist for a region the size of the continental United States, a huge number of 
references had to be reviewed. A task such as this is daunting given the breadth of publications on 
U.S. diatoms reports that have been recorded. It was decided to develop in concert with the check- 
list, a bibliography of the references consulted, and to include them with the taxon compilation. 

This current attempt could not be exhaustive, nor could it reconcile many of the discrepancies 
that are inherent in a preliminary checklist. It is hoped this listing provides a stepping stone for- 
ward to the development of a more accurate and, eventually, comprehensive understanding of the 
diatoms of inland environments of the United States. 



395 



396 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Methods 

The listing of diatom names was derived from those reports of inland habitats in the United 
States. Names are recorded where names were full (at least genus and species) and accepted by the 
author. Names with question marks next to them or designated with a single letter or number have 
not been included. A reference citing each taxon in the list is provided; a complete set of references 
for each name was not attempted. 

This listing of names reported in the literature, without original or cited interpretations means 
that there are redundancies in the names, either through taxonomic or nomenclatural synonymies. 
By nomenclatural (homotypic) synonymies, I mean there are different entries (names) based on a 
single type. Many of these occur in Fragilaria, Synedra, Achnanthes, Brachysira, Anomoeoneis, 
Navicula and others where there has been tremendous flux in the generic assignment of taxa. 
Despite the instability there has been in these names, much of this is well-known to those interest- 
ed in diatom taxonomy, and they are more easily traced due to the way authors are listed after the 
taxon names in botanical nomenclature. Taxonomic (heterotypic) synonymies, where more than 
one name, based on two (or more) types for what is the same diatom, are much harder to recog- 
nize, and actually very little of the monographic work necessary to discover these conditions has 
been done. An example would be: Gomphonema herculeanum var. robustum Grunow has been 
shown to be the same diatom as Gomphoneis herculeana (Ehrenberg) Cleve (Kociolek & Stoermer 
1988). 

Another situation that could result in a proliferation of names is typographic and/or ortho- 
graphic errors. Potential typographical errors were abundant in the literature consulted, and it is 
disconcerting to see how the same errors have been perpetuated from author to author. To reduce 
the number of entries due to these types of errors, I have, in some cases, made interpretations of 
these errors, especially in those situations when the context for the errors is more easy to interpret. 
For example, Loescher (1981, Table 1) Usts ''Achnanthes lanceolate (Breb.) Grun. var. lanceolata" 
in her prairie samples, and I have treated this as an orthographic error for Achnanthes lanceolata. 
Likewise, Burkholder & Wetzel (1989) list ''Epithemia adnata var. proboscidea (Kiitz.) Patr. 
comb, nov." (interpreted here as Epithemia adnata var. proboscidea) and ''Fragilaria cortonensis 
Kitt." (interpreted here as Fragilaria crotonensis), and as such I have not made separate entries for 
these orthographic errors. 

I have not tracked synonymies in either the traditions of VanLandingham's Catalogue (1967- 
1979) or Camburn & Charles (2000). In most of the cases, there has actually been very little orig- 
inal research to support the nomenclatural combinations made or perpetuated. Monographic or 
revisionary studies, (including consultation of types) are needed to support the numerous nomen- 
clatural changes made over the last decade or more. 

References included in the bibliography were from the published literature only; no unpub- 
lished manuscripts, reports, theses or dissertations were included. In some cases, especially in the 
case of research reports issued from programs or centers based at universities or local government 
agencies, publication was not easy to ascertain. Experimental works in situ are cited, but references 
to artificial systems are not. Published exsiccatae are included. Paleolimnological studies are, gen- 
erally, not included, except when surficial sediments were the focus of the study or report. 

The basis for much of the literature surveyed was the Maillard Library and Diatom Collection 
Library of the California Academy of Sciences, the reprint collections of Van Landingham, 
Rushforth, and Stoermer (now all at CAS) and the published exsiccatae catalogue of Edgar (1987). 



kociolek: freshwater diatoms of inland environments in the u.s. 397 

Results and Discussion 
Taxa 

A total of 167 genera and nearly 4500 names of diatoms at the level of species, variety or form 
are included in the list of diatoms (Section II: Species List of Diatoms of the United States). 
Naviciila has the largest number of names reported (over 900), followed by Pinnularia (340), 
Nitzschia (325), Gomphonema (237) and Eiinotia (223). Over forty genera are represented by a sin- 
gle name, and fifteen have only two names represented. Although many of these entries are nomen- 
clatural synonyms, it is interesting to note that many others represent newly discovered taxa, in the 
context of monographic revisions (Krammer 1992, 1997a, 1997b; 2002; Reichart 1999; Hamilton 
et al. 1995; Kociolek & Stoermer 1988), water quality studies (Potapova & Charles 2002, 2003; 
Morales 2003) and floristics (Stachura-Suchoples et al., accepted; Siver et al. 2005). 

Given the geogarphic size and complexity of the continental United States, the number of taxa 
in this compilation would appear modest compared to the check list developed for Great Britain 
(Hartley et al. 1986) and continental Europe (Krammer & Lange-Bertalot 1986, 1988, 1991a, 
1991b). That the U.S. flora has been developed almost exclusively with European taxon descrip- 
tions and keys (e.g., Hustedt 1927-1966, 1930), or with an incomplete flora (Patrick & Reimer 
1966, 1975) may have resulted in fewer taxa being reported from the country. 

References 

Over 1,200 citations are included in the Bibliography. There are over 150-1- years of publica- 
tions represented in this literature. The range of venues used to publish diatom reports includes 
those focused on phycology. botany, water quality assessment, state and local governments, feder- 
al government, water engineering, state academies, zoology, microscopy, ecology, hydrobiology, 
limnology and even speleology. These references show the importance of state and private acade- 
mies in the development of our knowledge of diatoms, especially the Academy of Natural Sciences 
of Philadelphia, and the Iowa Academy of Science. University centers have been Iowa State 
University, the University of Michigan, Bowling Green State University and Michigan State 
University, with faculty member at the latter three part of a lineage that has its roots at Iowa State 
and ANSP. Another important research center has been Brigham Young University. 

Some of the earliest reports on diatoms from U.S. freshwaters including sites from both coasts 
(e.g.. Ehrenberg 1854), and while there was tremendous development of our knowledge beyond 
these early works in the northeast, other places (like the deep South) have not yet developed sig- 
nificantly beyond the earliest reports. 

Areas that have been well-studied include the east coast, midwest, Great Lakes, Utah and 
Great Smoky Mountains National Park. Less studied areas include the southeastern U.S. (east, cen- 
tral and west), Pacific Northwest, California, and the Intermountain region. 

We are likely to associate investigation of water quality with the "green" period of the 1960s 
and 1970s, and there is a tremendous amount of literature relating to water quality issues from that 
period. However, there was significant attention and interest in the diatom communities of what we 
would now call eutrophic environments, especially as it related to sources of freshwater for human 
communities as far back as the late 19th century (e.g., Thomas & Chase 1887). 

Concluding Remarks 

The number of floristic studies of the U.S. freshwater flora is relatively few, given the size and 
diversity of the country. Recent floristic studies in some of the most relatively well-known areas, 
such as the east coast, continue to discover new taxa (Siver et al. 2005; Stachura-Suchoples et al. 



398 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

accepted), and monographic studies on some of the larger, more robust species in the genus 
Gomphoneis also identified many new taxa (Kociolek & Stoermer 1988). Thus, we still have a 
tremendous amount of work, floristic and monographic, to detail the diatoms of the U.S. Though 
the over 1 ,200 references included herein represent a large body of work, our knowledge of the 
flora has been constrained by the use of incomplete or European-based floras (Kociolek & 
Spaulding 2003). 

I believe the majority of new discoveries of taxa will not only come from relatively unexplored 
areas of the country (such as the description of a new, apparently endemic genus from playa lakes 
in Arizona-Spaulding et al. 2002), but also from oligotrophic habitats. An indication of the poten- 
tial for new discoveries from oligotrophic habitats comes from the work of Lange-Bertalot and 
Metzeltin (1996). They found in three oligotrophic lakes in Europe a total number of taxa equal to 
half of the total for all of Europe. Of the nearly 800 taxa found in these three oligotrophic lakes, 
over 125 were described as new or could not be assigned to known taxa. 

It is imperative to better understand the biodiversity of diatoms in the United States. The roles 
they play in providing ecosystem services (base of the food chain, producers of oxygen) and their 
apphcation to ecological problems, both theoretical and practical, provide strong rationales for a 
renewed commitment to develop the flora of the U.S. It is hoped this checklist, along with other 
tools, will facilitate work to the production of such a flora. 

Acknowledgments 

I am indebted in many ways to Gene Stoermer for his advice, guidance, and friendship. His 
insistence on high scientific standards has proven helpful beyond scientific endeavors. Dr. 
Elisabeth Fourtanier offered a constructive review of this manuscript as did Dr. Michele Aldrich. 

Literature Cited 

BoYER, C.S. 1927a. Synopsis of the North American Diatomaceae. Proceedings of the Academy of Natural 
Sciences of Philadelphia Supplement 78:1-228. 

BoYER, C.S. 1927b. Synopsis of the North American Diatomaceae. Proceeding of the Academy of Natural 
Sciences of Philadelphia Supplement 79:229-583. 

BURKHOLDER, J.M., AND R.G. Wetzel. 1989. Microbial colonization on natural and artificial macrophytes in 
a phosphorus-limited hardwater lake. Journal of Phycology 25:55-65. 

Camburn, K.E., AND D.F. Charles. 2000. Diatoms of Low Alkalinity Lakes in the Northeastern United States. 
Academy of Natural Sciences of Philadelphia, Special Publication 18. 152 pp. 

Edgar, R.K. 1978. The Jacob Whitman Bailey Diatom Collection at the Farlow Herbarium [of Harvard 
University]. Farlow Herbarium, Cambridge, Massachusetts, USA. iii + 155 pp. 

Ehrenberg, C.G 1854. Mikrogeologie. Das Erden und Felsen schaffende Wirken des unsichbar kleinen selb- 
ststdndigen Lebens aufder Erde. Leopold Voss, Leipzig, Germany. 374 pp. 

Fourtanier, E., and J.P. Kociolek. 1999. Catalogue of the diatom genera. Diatom Research 14:1-190. 

Fourtanier, E., and J.P. Kociolek. 2003. Addendum to the catalogue of diatom generic names. Diatom 
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Hamilton, P.B., M. Poulin, and D. Walker. 1995. Neidium hitchkockii (Ehrenberg) Cleve, a morphological- 
ly complex taxon within the genus Neidium (Naviculales, Bacillariophyta). Pages 6\-ll in J.P. Kociolek 
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Hartley, B., R. Ross, R., and D.M. Williams. 1986. A check-list of the freshwater, brackish and marine 
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HUSTEDT. F. 1927-1966. Die Kieselalgen Deutschlands, Osterreichs und der Schwiez mit Beriicksichtugung 

der iibrigen Lander Euwpas sowie der Angrenzenden Meeresgebiete. Rabenhorst Kryptogamenflora, 

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Germany. 
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Germany. 466p. 
KociOLEK. J. P. (submitted). Some thoughts on the development of a diatom flora for freshwater ecosystems 

in the continental United States and a listing of Recent taxa described from U.S. freshwaters. Proceedings 

of the California Academy of Sciences. 
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and R.G Sheath, eds.. Freshwater Algae of North America: Ecology and Classification, Chapt. 15. 

Academic Press, Boston, Massachusetts, USA. 
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Philadelphia 140:24-97. 
KR.A.MMER, K. 1992. Pinnularia. Fine Monographic der europaischen Taxa. Bibliotheca Diatomologica 

26:1-353. 
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Allgemeines und Encyonema Part. Bibliotheca Diatomologica 36:1-382. 
Krammer, K. 1997b. Die cymbelloiden Diatomeen. Fine Monographie der weltweit bekannten Taxa. Teil 2. 

Encyonema Part., Encyonopsis und Cymbellopsis. Bibliotheca Diatomologica 37:1^69. 
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Gantner, Verlag, Ruggell, Germany. 584 pp. 
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Frguanzungen zu Navicula (Lineolatae) und Gomphonema. In H. Fttl, G Gartner, J. Gerloff, H. Heynig, 

and D. Mollenhauer, eds., SUfiwasserflora von Mitteleuropa, 2/4. Gustav Fisher, Stuttgart, Germany. 437 

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Lange-Bertalot, H., and D. Metzeltin. 1996. Oligotrophie-Indikatoren. 800 Taxa reprasentativ fiir drei 

diverse Seen-Typen. Iconographia Diatomologica 2:1-390. 
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of the Iowa Academy of Science 88:63-69. 
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America, with comments on the taxonomy of the genus Synedra Fhrenberg. Proceedings of the Academy 

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400 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

tum/micropus, G. acuminatum sowie gomphonemoide Diatomeen aus dem Oberoligozan in Bohmen. 

Iconographia Diatomologica 8:1-203. 
SiVER, P.A., P.B. Hamilton, K. Stachura-Suchoples, and J.P. Kociolek. 2005. Freshwater diatom floras of 

North America: Cape Cod, Massachusetts, U.S.A. Diatomologica Iconographia 14:1-512. 
Spaulding, S.A., J.P. Kociolek, and D. Davis. 2002. A new diatom genus from a playa lake in New Mexico, 

USA with the description of two new species. European Journal of Phycology 37:135-143. 
Stachura-Suchoples, K., J.P. Kociolek, and PA. Siver. (In press.) A new Neidium species from Florida 

(U.S.A.) and comparison with A^. densestriatum (0strup) Krammer. Proceedings of the 17<^ International 

Diatom Symposium. 
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Their Synonyms. Vols. 1-8. J. Cramer, Lehre, Germany. 4654 pp. 

Section II: Species List of Diatoms of the United States 

Name Publication 

Acanthoceras zachariasii (Brun) Simonsen Stoermer et al. 1999 



Achnanthes acares Hohn & Hellerman Patrick & Reimer 

Achnanthes affinis Grunow Stoermer & Kreis 

Achnanthes altaica (Poretzsky) A. Cleve Stoermer et al. 



Achnanthes americana Cleve Boyer 1927b 



Achnanthes amoena Hustedt Stoermer & Kreis 

Achnanthes atacamae Hustedt Stoermer & Kreis 

Achnanthes austriaca Hustedt Cambum 

Achnanthes austriaca var. helvetica Hustedt Bateman & Rushforth 

Achnanthes biasolettiana (Kutzing) Grunow Stoermer & Kreis 



Achnanthes bicapitata Hustedt Cambum & Charles 2000 

Achnanthes biconfusa Van Landingham Cambum et al. 1978 

Achnanthes bioreti Germain Stoermer & Kreis 1978 

Achnanthes biporoma Hohn & Hellerman Hohn & Hellerman 1963 

Achnanthes bottnica (Cleve) Cleve Rushforth & Merkley 

Achnanthes brevipes C. Agardh Stoermer et al. 1999 

Achnanthes brevipes var. intermedia (Kiitzing) Cleve Stoermer et al. 1999 

Achnanthes calcar Cleve Stoermer & Kreis 1978 

Achnanthes chlidanos Hohn & Hellerman Cambum 1982 

Achnanthes chilensis var. subaequalis Reimer Whitford & Schumacher 1973 

Achnanthes clevei Gmnow Stoermer & Kreis 1978 

Achnanthes clevei var. rostrata Hustedt Stoermer & Kreis 1978 

Achnanthes coarctata (Brebisson) Gmnow Stoermer & Kreis 1978 

Achnanthes conspicua A. Mayer Stoermer & Kreis 1978 

Achnanthes conspicua var. brevistriata Hustedt Stoermer & Kreis 1978 

Achnanthes cotteriensis Foged Kaczmarska & Rushforth 1983 

Achnanthes crenulata Gmnow Rushforth & Merkley 

Achnanthes curvirostmm Bmn Patrick & Reimer 1966 

Achnanthes daui Foged Potapova & Charles 2003 

966 
978 
984 
978 
999 
978 
978 
978 
999 



Achnanthes decipiens Reimer Reimer 

Achnanthes deflexa Reimer Stoermer & Kreis 

Achnanthes deflexa var. alpestris Lowe & Kociolek Lowe & Kociolek 

Achnanthes delicatula (Kiitzing) Grunow Stoermer & Kreis 

Achnanthes depressa (Cleve) Hustedt Stoermer et al. 

Achnanthes detha Hohn & Hellerman Stoermer & Kreis 

Achnanthes didyma Hustedt Stoermer & Kreis 

Achnanthes dispar Cleve Stoermer & Kreis 

Achnanthes dispar var. angulata Hustedt Stoermer et al. 



966 
978 
999 



978 
978 
982 
984 
978 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 401 



Name Publication 



Achnanthes dispar var. fontellii A. Cleve Stoermer et al. 

Achnanthes duthii Screenivasa Stoermer et al. 

Achnanthes elUptica var. elongata A. Cleve Stoermer & Kreis 

Achnanthes exigua Gninow Stoermer & Kreis 

Achnanthes exigua var. constricta (Grunow) Hustedt Stoermer & Kreis 



Achnanthes exigua var. elliptica Hustedt Potapova & Charles 2003 

Achnanthes exigua var. heterovalva Krasske Stoermer & Kreis 1978 

Achnanthes exigua var. heterovalva f. semiaperta Guermeur Stoermer & Kreis 1978 

Achnanthes exilis Kutzing Stoermer & Kreis 1978 

Achnanthes flexella (Kiitzing) Brun Stoermer & Kreis 1978 

Achnanthes flexella var. alpestris Brun Stoermer et al. 1999 

Achnanthes gibberula Grunow Stoermer et al. 1999 

Achnanthes gracillima Hustedt Stoermer & Kreis 1978 

Achnanthes gracillina var. nipponica Sovereign Stoermer et al. 1999 

Achnanthes grana Hohn & Hellerman Patrick & Reimer 1966 

Achnanthes grimmei Krasske Patrick & Reimer 1966 

Achnanthes harveyi Reimer Cambum 1982 

Achnanthes hauckiana Grunow Stoermer & Kreis 1978 

Achnanthes hauckiana var. rostrata Schulz Stoermer & Kreis 1978 

Achnanthes helvetica (Hustedt) Lange-Bertalot Cambum & Charles 2000 



Achnanthes hettensis A. Cleve Stoermer & Kreis 

Achnanthes hudsonis Grunow in Van Heurck Patrick & Reimer 

Achnanthes hungarica (Grunow) Grunow Stoermer & Kreis 

Achnanthes hustedtii (Krasske) Reimer Patrick & Reimer 



Achnanthes imperfecta Schimanski Camburn & Charles 2000 



Achnanthes incognita Krasske Stoermer et al. 

Achnanthes inflata (Kutzing) Grunow Cambum 

Achnanthes inflata var. elata (Leud.-Fortm.) Hustedt Patrick & Reimer 

Achnanthes joursacense Heribaud Stoermer et al. 

Achnanthes kolbei Hustedt Stoermer et al. 

Achnanthes kryophila Peterson Stoermer & Kreis 

Achnanthes kryophila var. africana Cholnoky Stoermer & Kreis 

Achnanthes lanceolata (Brebisson) Grunow Stoermer & Kreis 

Achnanthes lanceolata var. abbreviata Reimer Hansmann 

Achnanthes lanceolata var. apiculata Patrick Stoermer & Kreis 

Achnanthes lanceolata var. bimaculata Hustedt Stoermer & Kreis 

Achnanthes lanceolata var. dubia Gmnow Stoermer & Kreis 

Achnanthes lanceolata var. elliptica Cleve Stoermer & Kreis 

Achnanthes lanceolata var. fossilis Patrick 

Achnanthes lanceolata var. genuina A. Mayer Stoermer & Kreis 

Achnanthes lanceolata var. haynaldii (Schaarsch.) Cleve Cambum & Lowe 

Achnanthes lanceolata var. lanceolatoides (Sovereign) Reimer Collins & Kalinsky 

Achnanthes lanceolata f. minor Schultz Whitford & Schumacher 

Achnanthes lanceolata var. omissa Reimer Stoermer & Kreis 

Achnanthes lanceolata var. robusta Hustedt Stoermer & Kreis 

Achnanthes lanceolata var. rhomboides A. Mayer Gaufin et al. 

Achnanthes lanceolata var. rostrata Hustedt Cambum 

Achnanthes lanceolata f. ventricosa Hustedt VanLandingham 

Achnanthes lanceolatoides Sovereign Stoermer & Kreis 

Achnanthes lapidosa Krasske Stoermer & Kreis 



Achnanthes lapidosa var. appalachiana (Cambum & Lowe) Lange-Bertalot Camburn & Charles 2000 



Achnanthes lapidosa var. lanceolata Patrick & Roberts 

Achnanthes lapponica (Hustedt) Hustedt Collins & Kalinsky 

Achnanthes lapponica var. fennica A. Cleve Clark & Rushforth 

Achnanthes lapponica var. ninckei (Guerm. & Manguin) Reimer Camburn 

Achnanthes laterostrata Hustedt Stoermer & Kreis 

Achnanthes lauenburgiana Hustedt Stoermer & Kreis 



999 
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966 
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966 



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402 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Name Publication 

Achnanthes lemmermannii Hustedt Stoermer & Kreis 1978 

Achnanthes levanderi Hustedt Stoermer & Kreis 1978 

Achnanthes levanderi var. helvetica Hustedt Stoermer et al. 1999 

Achnanthes lewisiana Patrick Stoermer & JCreis 1978 

Achnanthes linearis (W. Smith) Grunow Stoermer & Kreis 1978 

Achnanthes linearis var. pusilla Grunow Stoermer & Kreis 1978 

Achnanthes linearis f. curta H.L. Smith Stoermer & Kreis 1978 

Achnanthes linearis f. linearis Andresen & Stoermer 1978 

Achnanthes lutheri Hustedt Hansmann 1973 

Achnanthes macrocephala (Klitzing) Grunow Gaufin et al. 1976 

Achnanthes marginulata Grunow Stoermer & Kreis 1978 

Achnanthes microcephala (Klitzing) Grunow Stoermer & Kreis 1978 

Achnanthes minutissima Klitzing Stoermer & Kreis 1978 

Achnanthes minutissima vai'. cryptocephala Grunow Stoermer & Kreis 1978 

Achnanthes minutissima var. macrocephala Hustedt Hohn & Hellerman 1963 

Achnanthes minutissima var. robusta Hustedt Stoermer & Kreis 1978 

Achnanthes monela Hohn & Hellerman Hohn & Hellerman 1963 

Achnanthes montana Krasske Clark & Rushforth 1977 

Achnanthes nitiformis Lange-Bertalot Siver et al. 2005 

Achnanthes nodosa A. Cleve Stoermer & Kreis 1978 

Achnanthes nollii Bock Stoermer & Kreis 1978 

Achnanthes oblongella 0strup Cambum & Charles 2000 

Achnanthes oestrupi (A. Cleve) Hustedt Stoermer & Kreis 1978 

Achnanthes oestrupi var. lanceolata Hustedt Stoermer & Kreis 1978 

Achnanthes oestrupii var. parvula Patrick Patrick 1945 

Achnanthes oestrupii var. pungens (A. Cleve) Lange-Bertalot Stoermer et al. 1999 

Achnanthes orientalis Petit Rushforth & Merkley 1988 

Achnanthes pachypus Montague Patrick & Reimer 1966 

Achnanthes peragalli Brun & Heribaud Stoermer & Kreis 1978 

Achnanthes peragalli var. fossilis Tempere & Peragallo Stoermer & Kreis 1978 

Achnanthes peragalli var. parvula (Patrick) Reimer Stoermer & Kreis 1978 

Achnanthes pericava Carter Gaiser & Johansen 2000 

Achnanthes pinnata Hustedt Stoermer & Kreis 1978 

Achnanthes ploenensis Hustedt Stoermer & Kreis 1978 

Achnanthes prava Sovereign Patrick & Reimer 1966 

Achnanthes procera Hustedt Stoermer & Kreis 1978 

Achnanthes pseudolinearis Hustedt Cambum 1982 

Achnanthes pseudoobliqua Simonsen Kalinsky 1983 

Achnanthes pseudotanensis A. Cleve Patrick & Reimer 1966 

Achnanthes pusilla Grunow in Cleve & Grunow Hamilton et al. 1992 

Achnanthes quadratarea (0strup) Moller ex Foged Hamilton et al. 1992 

Achnanthes recurvata Hustedt Stoermer et al. 1999 

Achnanthes reimeri Cambum Cambum 1982 

Achnanthes ricula Hohn & Hellerman Hohn & Hellerman 1963 

Achnanthes rosenstockii Lange-Bertalot Cambum & Charles 2000 

Achnanthes rostrata 0stmp Hamilton et al. 1992 

Achnanthes rapestoides Hohn Patrick & Reimer 1966 

Achnanthes saxonica Krasske Patrick & Reimer 1966 

Achnanthes simplex Hustedt Kaczmarska & Rushforth 1983 

Achnanthes stewartii Patrick Cambum 1982 

Achnanthes subatomus Hustedt Hohn & Hellerman 1963 

Achnanthes subhudsonis var. kraeuselii Cholnoky Hohn & Hellerman 1963 

Achnanthes sublaevis Hustedt Stoermer & Kreis 1978 

Achnanthes sublaevis var. crassa Reimer Cambum 1982 

Achnanthes submarina Hustedt Kalinsky 1983 

Achnanthes subrostrata Hustedt Dodd 1987 

Achnanthes subrostrata var. appalachiana Cambum & Lowe Cambum 1982 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 403 



Name Publication 

Achnanthes subsaloides Hustedt Stoermer & Kreis 1978 

Achnanthes subsalsa Petersen 

Achnanthes subsessilis Kutzing KaUnsky 1983 

Achnanthes suchlandtii Hustedt Stoermer & Kreis 1978 

Achnanthes taeniata Grunovv in Cleve & Grunow Patrick & Roberts 1979 

Achnanthes temperei Peragallo in Tempere & Peragallo Boyer 1927b 

Achnanthes thermalis (Rabenhorst) Schonfeld Stoermer et al. 1999 

Achnanthes trinodis (W. Smith) Grunow Stoermer & Kreis 1978 

Achnanthes tropica Hustedt Kaczmarska & Rushforth 1983 

Achnanthes ventralis (Krasske) Lange-Bertalot Stoermer et al. 1999 

.A.chnanthes ventraloconfusa f. simplex Lange-Bertalot Stoermer et al. 1999 

Achnanthes ventricosa Ehrenberg Tempere & Peragallo 1909 

Achnanthes wellsiae Reimer Stoermer & Kreis 1978 

Achnanthidium affme (Grunow) Czamecki Stoermer et al. 1999 

Achnanthidium alpestre (Lowe & Kociolek) Lowe & Kociolek in Johansen et al Potapova & Ponader 2004 

Achnanthidium bioreti (Germain) Edlund Stoermer et al. 1999 

Achnanthidium brevipes var. intermedia (Kutzing) Cleve Stoermer et al. 1999 

Achnanthidium clevei (Grunow in Cleve & Grunow) Czarnecki Stoermer et al. 1999 

Achnanthidium clevei var. rostratum (Hustedt) Edlund Stoermer et al. 1999 

Achnanthidium crassum (Hustedt) Potapova & Ponader Potapova & Ponader 2004 

Achnanthidium deflexum (Reimer) Kingston Potapova & Charles 2003 

Achnanthidium delicatulum Kiitzing Stoermer et al. 1999 

Achnanthidium duthii (Sreen.) Edlund Stoermer et al. 1999 

Achnanthidium exiguum (Grunow) Czamecki Stoermer et al. 1999 

Achnanthidium exiguum var. constrictum (Grunow) Andresen et al Andresen et al. 2000 

Achnanthidium exiguum var. heterovalvum (Krasske) Czarnecki Stoermer et al. 1999 

Achnanthidium exilis (Kiitzing) Bukhtiyarova Stoermer et al. 1999 

Achnanthidium flexellum (Kutzing) Brebisson Stoermer & Kreis 1978 

Achnanthidium hauckianum (Grunow) Czamecki Stoermer et al. 1999 

Achnanthidium hungaricum Grunow Stoermer et al. 1999 

Achnanthidium kryophila (Petersen) Bukhtiyarova Stoermer et al. 1999 

Achnanthidium lanceolatum Brebisson in Kutzing Stoermer et al. 1999 

Achnanthidium lanceolatum var. elliptica Cleve Stoermer et al. 1999 

Achnanthidium lanceolatum var. genuinum Mayer Stoermer et al. 1999 

Achnanthidium lanceolata var. haynaldii (Schaarsch.) Cleve Stoermer et al. 1999 

Achnanthidium lapponicum Hustedt Hohn 195 1 

Achnanthidium levanderi (Hustedt) Czarnecki Stoermer et al. 1999 

Achnanthidium lineare W. Smith Stoermer et al. 1999 

Achnanthidium microcephalum (Kutzing) vide Rabenhorst Stoermer et al. 1999 

Achnanthidium minutissimum (Kutzing) Czarnecki Stoermer et al. 1999 

Achnanthidium peragalli Bmn & Heribaud in Heribaud Stoermer et al. 1999 

Achnanthidium rivulare Potapova & Ponader Potapova & Ponader 2004 

Achnanthidium semiaperta (Guermeur) Andresen et al Andresen et al. 2000 

Achnanthidium trinodis Amott ex Ralfs in Pritchard Stoermer et al. 1999 

Actinella punctata Lewis Boyer 1927a 

Actinocyclus australis Gmnow Tempere & Peragallo 1913 

Actinocyclus barkelyi Grunow Tempere & Peragallo 1913 

Actinocyclus niagarae H.L. Smith Stoermer & Kreis 1978 

Actinocyclus normanii (Gregory) Hustedt Stoermer et al. 1999 

Actinocyclus normanii f. subsalsa (Juhl.-Dannf.) Hustedt Stoermer et al. 1999 

Actinoptychus sp Stoermer & Kreis 1978 

Adlafia suchlandtii (Hustedt^ Lange-Bertalot in Moser et al Johansen et al. 2004 



404 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Name Publication 

Amphicampa eruca Ehrenberg Boyer 1927a 

Amphicampa hemicyclus (Ehrenberg) Karsten Rushforth & Merkley 

Amphicampa mirabihs Ehrenberg ex Ralfs Patrick & Reimer 

Amphipleura artica Patrick & Freese Stoermer & Kreis 

Amphipleura kriegeriana (Krasske) Hustedt Stoermer et al. 

Amphipleura lindheimeri Grunow Patrick & Reimer 

Amphipleura pellucida (Kutzing) Kutzing Stoermer & Kreis 

Amphipleura rutilans (Trentepohl) Cleve Rushforth & Merkley 

Amphipleura sigmoidea W. Smith Stoermer & Kreis 

Amphiprora alata (Ehrenberg) Kutzing Stoermer & Kreis 

Amphiprora calumetica Thomas Stoermer & Kreis 

Amphiprora conspicua Greville Patrick & Reimer 

Amphiprora costata Hustedt Whitford & Schumacher 

Amphiprora navicularis Ehrenberg Ehrenberg 

Amphiprora nereis Lewis Patrick & Reimer 

Amphiprora omata J.W. Bailey Stoermer & Kreis 

Amphiprora paludosa W. Smith Stoermer & Kreis 

Amphiprora pulchra J.W. Bailey J.W. Bailey 

Amphitropis omata (J.W. Bailey) Grunow Stoermer & Kreis 

Amphitropis paludosa var. duplex Donkin Cleve & Moller 

Amphitropis paludosa var. pokomiana Grunow Cleve & Moller 

Amphora acutiuscula Kutzing Patrick & Reimer 

Amphora amphioxys J.W. Bailey Ehrenberg 

Amphora angularis Gregory Kalinsky 

Amphora angusta Gregory Patrick & Reimer 

Amphora angusta var. oblongella Grunow Kalinsky 

Amphora angusta var. ventricosa (Gregory) Cleve Kalinsky 

Amphora aponina Kutzing Patrick & Reimer 

Amphora arcus var. sulcata (A. Smith) Cleve Kaczmarska & Rushforth 

Amphora birugula Hohn Cambum 

Amphora bullatoides Hohn & Hellerman Stoermer & Kreis 

Amphora calumetica (Thomas) M. Peragallo Stoermer & Kreis 

Amphora caroliniana Giffen Kalinsky 

Amphora cistula var. maculata (Kutzing) Van Heurck Prcscott & Dillard 

Amphora clevei Grunow in A. Schmidt Patrick & Reimer 

Amphora coffeaeformis (Agardh) Kutzing Stoermer & Kreis 

Amphora coffeaeformis var. acutiuscula (Kutzing) Kiitzing Stoermer & Kreis 

Amphora coffeaeformis var. fossilis Pantocsek Kaczmarska & Rushforth 

Amphora coffeaeformis var. perpusilla Grunow Kaczmarska & Rushforth 

Amphora coffeiformis var. salinarum Grunow Patrick & Reimer 

Amphora cognata Cholnoky Rushforth & Merkley 

Amphora commutata Grunow Stoermer & Kreis 

Amphora crassa Gregory Rushforth & Merkley 

Amphora crucifera A. Cleve Stoermer et al. 

Amphora cruciferoides Stoermer & Yang Stoermer & Kreis 

Amphora cymbifera Gregory' Patrick & Reimer 

Amphora delicatissima Krasske Stoermer & Kreis 

Amphora delphina L. W. Bailey Boyer 1927b 

Amphora delphinea var. minor Tempere & Peragallo 

Amphora exigua Gregory Stoermer & Kreis 

Amphora fonticola Maillard Stoermer & Kreis 

Amphora fontinalis Hustedt Cambum 

Amphora fusca A. Schmidt Gaiser & Johansen 2000 

Amphora gigas Ehrenberg Stoermer & Kreis 1978 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 405 



Name Publication 

Amphora gracilis Ehrenberg 

Amphora granulata Gregor}' Kalinsky 

Amphora hemicycla Stoermer & Yang Stoermer & Kreis 

.\mphora holsatica Hustedt Patricic & Reimer 

Amphora huronensis Stoermer & Yang Stoermer & Kreis 

Amphora hyalina Kiitzing Kaczmarska & Rushforth 

Amphora inariensis Krammer Stoermer et al. 

Amphora libyca Ehrenberg Stoermer & Kreis 

Amphora hneata Gregory Patrick & Reimer 

Amphora lineolata (Ehrenberg) Ehrenberg Prescott & Dillard 

.Amphora macilenta Gregory Kaczmarska & Rushforth 

Amphora marina (W. Smith) Van Heurck Rushforth & Merldey 

.Amphora menisca Hohn & Hellerman Hohn & Hellerman 

Amphora michiganensis Stoermer & Yang Stoermer & Kreis 

Amphora montana Krasske Stoermer & Kreis 

.\mphora neglecta Stoermer & Yang Stoermer & Kreis 

Amphora normanii Rabenhorst Stoermer & Kreis 

Amphora ocellata Ehrenberg 

Amphora ohgotraphenta Lange-Bertalot Stoermer et al. 

Amphora ovalis (Kiitzing) Kiitzing Stoermer & Kreis 

Amphora ovalis var. affmis (Kutzing) Van Heurck ex De Toni Cambum 

Amphora ovalis var. constricta Skvortzow Stoermer & Kreis 

Amphora ovalis var. gracilis (Ehrenberg) Van Heurck Stoermer & Kreis 

Amphora ovalis var. libyca (Ehrenberg) Cleve Stoermer & Kreis 

Amphora ovalis var. minor H.H. Chase Stoermer & Kreis 

Amphora ovalis var. minutissima (W. Smith) Hurler Stoermer & Kreis 

Amphora ovalis var. pediculus (Kutzing). Van Heurck Stoermer & Kreis 

Amphora parallelistriata Manguin Stoermer & Kreis 

Amphora parallelistriata var. manguinii Carter Clark & Rushforth 

Amphora pediculus var. minor Grunow Whitford & Schumacher 

Amphora pellucida Gregory Patrick & Reimer 

Amphora perpusilla (Grunow) Grunow Stoermer & Kreis 

Amphora proteus Gregory Patrick & Reimer 

Amphora proteus var. oculata Peragallo Kaczmarska & Rushforth 

Amphora rimosa Ehrenberg Patrick & Reimer 

Amphora rotunda Skvortzow Stoermer & Kreis 

Amphora sabiniana Reimer Patrick & Reimer 

Amphora salina W. Smith Elmore 

Amphora sibirica Skvortzow in Skvortzow & Meyer Stoermer & Kreis 

Amphora subcostulata Stoermer & Yang Stoermer & Kreis 

Amphora submontana Hustedt Cambum 

Amphora tenerrima Hustedt Kalinsky 

Amphora tenuissima Hustedt Kaczmarska & Rushforth 

Amphora tenuistriata Manguin Hohn & Hellerman 

Amphora thermalis Hustedt Drum 

Amphora thumensis (Mayer) A. Cleve Stoermer et al. 

Amphora turgida Gregory Patrick & Reimer 

Amphora veneta Kutzing Stoermer & Kreis 

Amphora veneta var. angularis Stoermer & Kreis 

Amphora veneta var. capitata Haworth Stoermer & Kreis 

Amphora ventricosa Gregory Kalinsky 

Aneumastus carolinianus (Patrick; Lange-Bertalot Lange-Bertalot 2001 

Aneumastus minor (Hustedt) Lange-Bertalot Stoermer et al. 1999 

Aneuma.stus stroesei (0strupj Mann in Round et al Stoermer et al. 1999 

Aneumastus tusculus (Ehrenberg) Mann in Round et al Stoermer et al. 1999 

Aneumastus tusculus f. minor (Hustedt) Andresen et al Andresen et al. 2000 



406 



PROCEEDINGS OF THE CALIFORNIA AC ADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 



Name 



Publication 



Aneumastus tusculus f. obtusum (Hustedt) Andresen et al Andresen et al. 2000 

Aneumastus tusculus var. rostratus (Hustedt) Andresen et al Andresen et al. 2000 



Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 
Anomoeone 



s brachysira (Brebisson) Grunow Cambum & Charles 2000 

s brachysira var. zellensis (Grunow) Krammer Cambum & Charles 2000 

s costata (Kiitzing) Hustedt Stoermer & Kreis 1978 

s exilis Kiitzing Stoermer & Kreis 1978 

s exilis var. lanceolata A. Mayer Bateman & Rushforth 1984 

s exilis var. thermalis (Grunow) Cleve Patrick & Reimer 1966 

s fogedii Reimer Reimer 1982 

s follis (Ehrenberg) Cleve Stoermer & Kreis 1978 

s follis var. hannae Reimer Patrick & Reimer 1966 

s follis var. fossilis Reimer Patrick & Reimer 1966 

s paludigena Scherer Scherer 1988 

s polygramma (Ehrenberg) Cleve Rushforth & Merkley 1988 

s polygramma var. platensis Frenguelli Patrick & Reimer 1966 

s sculpta (Ehrenberg) O. Mtiller Stoermer & Kreis 1978 

s sculpta var. major Cleve Patrick & Reimer 1966 

s serians Brebisson Stoermer & Kreis 1978 

s serians f. undulata Hustedt Hustedt 1959 

s serians var. acuta Hustedt Hansmann 1973 

s serians var. apiculata (Lewis) Boyer Boyer 1927b 

s serians var. brachysira (Brebisson) Hustedt Stoermer & Kreis 1978 

s serians var. brachysira f. thermalis (Grunow) Hustedt Sovereign 1958 

s serians var. minor Grunow Boyer 1927b 

s sphaerophora (Ehrenberg) Pfitzer Stoermer & Kreis 1978 

s sphaerophora var. biceps Ehrenberg Ehrenberg 1856 

s sphaerophora f. costata (Kiitzing) A.M. Schmid Stoermer et al. 1999 

s sphaerophora var. guentheri O. Miiller Patrick & Reimer 1966 

s sphaerophora var. minor Kociolek & Herbst Kociolek & Herbst 1992 

s sphaerophora var. sculpta (Ehrenberg) O. Mtiller Stoermer & Kreis 1978 

s styriaca (Grunow) Hustedt Stoermer & Kreis 1978 

s variabilis (Ross) Reimer Reimer 1966 

s vitrea (Grunow) Ross Stoermer & Kreis 1978 

s vitrea f. lanceolata (A. Mayer) Mogh Gaufin et al. 1976 

s vitrea var. gomphonemacea (Grunow) Mogh Gaufin et al. 1976 

s zellensis (Grunow) Cleve Stoermer & Kreis 1978 

s zellensis f. difficilis (Grunow in Van Heurck) Hustedt Stoermer et al. 1999 



Anorthoneis dulcis Hein Hein 1991 

Asterionella bleakleyi W. Smith Stoermer & Kreis 1978 

Asterionella formosa Hassall Stoermer & Kreis 1978 

Asterionella formosa var. acaroides Lemmerman Stoermer & Kreis 1978 

Asterionella formosa var. gracillima (Hantzsch) Grunow in Van Heurck Stoermer & Kreis 1978 

Asterionella formosa var. subtilis Grunow Stoermer & Kreis 1978 

Asterionella formosa var. subtilissima Grunow Starrett & Patrick 1952 

Asterionella gracillima (Hantzsch) Heiberg Stoermer & Kreis 1978 

Asterionella ralfsii W. Smith Patrick 1945 

Asterionella ralfsii var. americana Komer Dixit & Smol 1995 

Attheva zachariasi Brun Stoermer & Kreis 1978 



Aulacoseira agassizii (Ostenfeld) Simonsen Stoermer et al. 1999 

Aulacoseira agassizii var. malayensis (Hustedt) Simonsen Stoermer et al. 1999 

Aulacoseira alpigena (Grunow) Krammer Stoermer et al. 1999 

Aulacoseira ambigua (Grunow) Simonsen Stoermer et al. 1999 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 407 



Name Publication 

Aulacoseira canadensis (Hustedt) Simonsen Stoermer et al. 1999 

Aulacoseira crassipunctata Krammer Cambum & Charles 2000 

Aulacoseira distans (Ehrenberg) Simonsen Stoermer et al. 1999 

Aulacoseira distans var. africana (O. Muller) Simonsen Hamilton et al. 1992 

Aulacoseira distans var. alpigena (Ehrenberg) Simonsen Cambum 1982 

Aulacoseira distans var. limnetica (O. Muller) Simonsen Stoermer et al. 1999 

Aulacoseira distans var. nivalis (W. Smith) Haworth Cambum & Charles 2000 

Aulacoseira distans var. nivaloides (Cambum) Haworth Hamilton et al. 1992 

Aulacoseira distans var. septentrionalis Cambum & Charles Cambum & Charles 2000 

Aulacoseira distans var. tenella (Nygaard) Ross in Hartley Hamilton et al. 1992 

Aulacoseira granulata (Ehrenberg) Simonsen Cambum 1982 

Aulacoseira granulata var. angustissima (O. Muller) Simonsen Cambum 1982 

Aulacoseira granulata var. angustissima f. spiralis (Hustedt) Czamecki & Reinke Stoermer et al. 1999 

.A.ulacoseira granulata var. muzzanensis (Meister) Simonsen Stoermer et al. 1999 

Aulacoseira herzogii Lemmermann Hickel & Hakansson 1991 

Aulacoseira humilis (Cleve-Euler) Simonsen Cambum & Charles 2000 

.Aulacoseira islandica (O. Miiller) Simonsen Stoermer et al. 1999 

.Aulacoseira islandica subsp. helvetica (O. Muller) Simonsen Stoermer et al. 1999 

Aulacoseira italica (Ehrenberg) Simonsen Stoermer et al. 1999 

.Aulacoseira italica var. tenuissima (Grunow) Simonsen Stoermer et al. 1999 

.Aulacoseira italica var. valida (Gmnow) Simonsen Stoermer et al. 1999 

Aulacoseira italica subsp. subarctica (O. Muller) Simonsen Stoermer et al. 1999 

Aulacoseira lacustris (Grunow) Krammer Camburn & Charles 2000 

Aulacoseira lirata (Ehrenberg) Kiitzing Hamilton et al. 1992 

.Aulacoseira lirata var. biseriata (Gmnow) Haworth Hamilton et al. 1992 

Aulacoseira lirata var. lacustris (Grunow) Ross in Haartley Hamilton et al. 1992 

Aulacoseira lirata var. alpigena (Gmnow) Haworth Stoermer et al. 1999 

Aulacoseira lirata var. perglabra (0stmp) Ross in Hartley Hamilton et al. 1992 

Aulacoseira lirata var. perglabra f. florinae (Cambum) Haworth Hamilton et al. 1992 

Aulacoseira nygaardii (Cambum) Cambum in Cambum & Charles Hamilton et al. 1992 

Aulacoseira perglabra (0stmp) Haworth Cambum & Charles 2000 

Aulacoseira perglabra var. floriniae (Cambum) Haworth Cambum & Charles 2000 

Aulacoseira subarctica (O. Miiller) Haworth Stoermer et al. 1999 

Aulacoseira tenella (Nygaard) Simonsen Cambum & Charles 2000 

Aulacoseira valida (Gmnow) Krammer Stoermer et al. 1999 

Bacillaria furcata Ehrenberg 1856 

Bacillaria cuneata Ehrenberg 1856 

Bacillaria paradoxa Gmelin Stoermer & Kreis 1978 

Bacillaria paradoxa var. Uimidula Gmnow Collins & Kalinsky 1977 

Bacillaria paxillifer (O.F. Muller) Hendey Stoermer & Kreis 1978 

Bacillaria vulgaris Ehrenberg 1856 

Biddulphia laevis Ehrenberg Cambum 1982 

Biddulphia polymorpha Ehrenberg Whitford 1956 

Biremis circumtexta (Meister) Lange-Bertalot & Witkowski Potapova & Charles 2003 

Biremis undulata (Schulz) Andresen et al Edlund et al. 2001 

Biremis zachariasi CReichelt) Edlund et al Edlund et al. 2001 

Brachysira aponina Kiitzing Kaczmarska & Rushforth 1983 

Brachysira arctoborealis Wolfe & Kling Wolfe & Kling 2001 

Brachysira brebissonii Ross Potapova & Charles 2003 

Brachysira exilis CKutzingj Round & D.G Mann Stoermer et al. 1999 

Brachysira exilis f. undulata Kisselev Kaczmarska & Rushforth 1983 

Brachysira follis fEhrenbergj R. Ross in Hartley Stoermer et al. 1999 

Brachysira gravida Shayler & Siver Shayler & Siver 2004 



408 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Name Publication 

Brachysira brebissonii R. Ross in Hartley Stoermer et al. 1999 

Brachysira microcephala (Grunow) Compere Potapova & Charles 2003 

Brachysira neoacuta Lange-Bertalot Gaiser & Johansen 2000 

Brachysira neoexilis Lange-Bertalot Potapova & Charles 2002 

Brachysira serians (Brebisson ex Kiitzing) Round & D.G Mann Stoermer et al. 1999 

Brachysira serians var. acuta (Hustedt) Hamilton in Hamilton et al Hamilton et al. 1992 

Brachysira sphaerophora (Kiitzing) Round & D.G. Mann Stoermer et al. 1999 

Brachysira styriaca (Grunow in Van Heurck) R. Ross in Hartley Stoermer et al. 1999 

Brachysira vitrea (Grunow) R. Ross in Hartley Stoermer et al. 1999 

Brachysira zellensis (Grunow) Round & D.G. Mann Stoermer et al. 1999 

Brachysira zellensis f. difFicilis (Grunow in Van Heurck) Hamilton in Hamilton et al Stoermer et al. 1999 

Brebissonia boeckii (Ehrenberg) Grunow Boyer 1927b 

Brebissonia interposita (Lewis) Kuntze Elmore 1922 

Brebissonia palmeri Boyer Smith 1950 

Brebissonia vulgaris (Thwaites) Van Heurck Elmore 1922 



Calone: 
Calone: 
Calone: 
Calone; 
Calone 
Calone 
Calone: 
Calone: 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone: 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 
Calone 



s aequatorialis Hustedt Dodd 1987 

s aerophila Bock Collins & Kalinsky 1977 

s alpestris (Grunow) Cleve Stoermer & Kreis 1978 

s amphisbaena (Bory) Cleve Stoermer & Kreis 1978 

s amphisbaena var. subsalina (Donkin) Cleve Hohn 1951 

s bacillaris (Gregory) Cleve Stoermer & Kreis 1978 

s bacillaris var. thermalis (Grunow) A. Cleve Stoermer & Kreis 1978 

s bacillum (Grunow) Cleve Stoermer & Kreis 1978 

s bacillum var. angusta A. Mayer Dodd 1987 

s bacillum f. fonticola (Grunow) A. Mayer Dodd 1987 

s bacillum var. fontinalis Hustedt Stoermer & Kreis 1978 

s bacillum var. lancettula (Schulz) Hustedt Stoermer & Kreis 1978 

s backmanii A. Cleve Czamecki & Blinn 1978 

s clevei (Lagesterdt) Cleve Stoermer & Kreis 1978 

s clevei var. undulata Krasske Stoermer et al. 1999 

s columbiensis Cleve Boyer 1927b 

s fasciata (lagestedt) Cleve Boyer 1927b 

s fenzlii (Grunow) Patrick Patrick & Reimer 1966 

s fenzlioides Cleve-Euler Rushforth & Squires 1985 

s fontinalis (Grunow) Kange-Bertalot & Reichardt Stoermer et al. 1999 

s formosa Gregory Stoermer & Kreis 1978 

s hebes (Ralfs) Patrick Patrick & Reimer 1966 

s holstii Cleve Cleve 1894 

s hultenii Petersen Dodd 1987 

s hyalina Hustedt Stoermer & Kreis 1978 

s lagerstedtii Cholnoky Dodd 1987 

s lamella Zakrzewski Collins & Kalinsky 1977 

s lancettula (Schulz) Lange-Bertalot & Witkowski Stoermer et al. 1999 

s latiuscula var. reimeri Czamecki & Blinn Czamecki 1979 

s lenzii Krasske Dodd 1987 

s leptosoma (Gmnow) Krammer Stoermer et al. 1999 

s lewisii Patrick Stoermer & Kreis 1978 

s lewisii var. inflata (Schultze) Patrick Patrick & Reimer 1966 

s liber (W. Smith) Cleve Stoermer et al. 1999 

s limosa (Kutzing) Patrick Stoermer & Kreis 1978 

s limosa var. gibberala (Kutzing) Gmnow Stoermer et al. 1999 

s limosa var. subinflata Gmnow Stoermer et al. 1999 

s limosa var. undulata Gmnow Stoermer et al. 1999 

s molaris (Gmnow) Krammer Stoermer et al. 1999 

s nubicola (Gmnow) Cleve Stoermer & Kreis 1978 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 409 



Name Publication 

Caloneis obtusa (W. Smith) Cleve Boyer 1927b 

Caloneis oregonica (Ehrenberg) Patrick Grimes & Rushforth 1982 

Caloneis permagna (Bailey) Cleve Rushforth & Squires 1985 

Caloneis permagna var. lewisiana Boyer Starrett & Patrick 1952 

Caloneis pseudoclevei Cholnoky Patrick & Reimer 1966 

Caloneis pseudoschummaniana (Hustedt) Simonsen Simonsen 1987 

Caloneis pulchra Messik Stoermer & Kreis 1978 

Caloneis pulchra var. interrupta Gandhi Stoermer & Kreis 1978 

Caloneis salebrastriata Hohn Patrick & Reimer 1966 

Caloneis schumanniana (Grunow) Cleve Stoermer & Kreis 1978 

Caloneis schumanniana var. biconstricta Grunow Stoermer & Kreis 1978 

Caloneis schumanniana var. biconstricta f. baikalensis Skv Johansen et al. 2004 

Caloneis schumanniana var. fasciata Hustedt Grimes & Rushforth 1982 

Caloneis schumanniana var. lancettula Hustedt Stoermer et al. 1999 

Caloneis schumanniana var. linearis Hustedt Grimes & Rushforth 1982 

Caloneis silicula (Ehrenberg) Cleve Stoermer & Kreis 1978 

Caloneis sihcula var. alpina Cleve Patrick 1945 

Caloneis silicula var. brevistriata O. Miiller Czarnecki & Blinn 1978 

Caloneis sihcula var. gibberula (Kiitzing) Cleve Boyer 1927b 

Caloneis silicula var. inflata (Grunow) Cleve Boyer 1927b 

Caloneis sihcula var. minuta (Grunow) Cleve Patrick 1961 

Caloneis silicula var. truncatula (Grunow) Meister Stoermer & Kreis 1978 

Caloneis silicula var. limosa Van Landingham Stoermer et al. 1999 

Caloneis silicula var. tumida Hustedt Czarnecki & Blinn 1978 

Caloneis silicula var. undulata (Grunow) Cleve Stoermer et al. 1999 

Caloneis silicula var. ventricosa Patrick 1968 

Caloneis speciosa (Hustedt) Boyer Boyer 1927b 

Caloneis tenuis (Gregory) Krammer Stoermer et al. 1999 

Caloneis trinodis (Lewis) Boyer Boyer 1927b 

Caloneis trinodis var. inflata Schultze Patrick & Reimer 1966 

Caloneis undosa Krammer Camburn & Charles 2000 

Caloneis undulata (Gregory) Krammer Stoermer et al. 1999 

Caloneis ventricosa (Ehrenberg) Meister Stoermer & Kreis 1978 

Caloneis ventricosa var. alpina Patrick Collins & Kalinsky 1977 

Caloneis ventricosa var. inflata Collins & Kalinsky 1977 

Caloneis ventricosa var. minuta (Grunow) Patrick Stoermer & Kreis 1978 

Caloneis ventricosa var. truncatula (Grunow) Meister Stoermer & Kreis 1978 

Caloneis ventricosa var. subundulata (Grunow) Patrick Patrick & Reimer 1966 

Caloneis westii (W. Smith) Hendey Kaczmarska & Rushforth 1983 

Caloneis zachariasii Reichelt Prescott & Dillard 1979 

Campylodiscus alaetus Setty Rushforth & Merkley 1988 

Campylodiscus american Ehrenberg 1856 

Campylodiscus bicostatus W. Smith Elmore 1922 

Campylodiscus clypeus Ehrenberg Stoermer et al. 1999 

Campylodiscus costatus W. Smith Tempere & Peragallo 1909 

Campylodiscus cribrosus W. Smith Stoermer & Kreis 1978 

Campylodiscus decorus Brebisson Stoermer & Kreis 1978 

Campylodiscus echensis Ehrenberg Stoermer et al. 1999 

Campylodiscus ehrenbergii Ralfs Rushforth & Merkley 1988 

Campylodiscus eiowanus Ehrenberg 1856 

Campylodiscus hibemicus Ehrenberg Stoermer & Kreis 1978 

Campylodiscus noricus Ehrenberg Stoermer & Kreis 1978 

Campylodiscus noricus var. hibemica (Ehrenberg) Grunow Stoermer & Kreis 1978 

Campylodiscus spiralis Myers 1898 

Capartogramma crucicula (Grunow) Ross Stoermer & Kreis 1978 



410 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Name Publication 

Catacombus gaillonii (Bory) Williams & Round Stoermer et al. 1999 

Cavinula cocconeiformis (Gregory ex Greville) D.G. Mann & Stickle in Round et al Stoermer et al. 1999 

Cavinula intractata (Hustedt) Lange-Bertalot Stoermer et al. 1999 

Cavinula jaemefeltii (Hustedt) D.G. Mann & Stickle in Round et al Stoermer et al. 1999 

Cavinula lacustris (Gregory) D.G. Mann & Stickle in Round et al Stoermer et al. 1999 

Cavinula pseudoscutiformis (Hustedt) D.G. Mann & Stickle in Round et al Stoermer et al. 1999 

Cavinula scutelloides (W. Smith) Lange-Bertalot Stoermer et al. 1999 

Cavinula scutiformis (Grunow ex A. Schmidt) D.G. Mann & Stickle in Round et al Stoermer et al. 1999 

Cavinula variostriata (Krasske) D.G. Mann & Stickle in Round et al Stoermer et al. 1999 

Ceratoneis arcus Kiitzing Stoermer & Kreis 1978 

Ceratoneis arcus van amphioxys (Rabenhorst) Grunow Stoermer & Kreis 1978 

Ceratoneis fasciolata Ehrenberg 1856 

Chamaepinnularia begeri (Krasske) Lange-Bertalot Stoermer et al. 1999 

Chamaepinnularia bremensis (Hustedt) Lange-Bertalot in Lange-Bertalot & Metzeltin Siver et al. 2005 

Chamaepinnularia evanida Lange-Bertalot Stoermer et al. 1999 

Chamaepinnularia margaritacea (Hustedt) Lange-Bertalot Johansen et al. 2004 

Chamaepinnularia mediocris (Krasske) Lange-Bertalot Stoermer et al. 1999 

Chamaepinnularia soehrensis (Krasske) Lange-Bertalot Stoermer et al. 1999 

Chamaepinnularia soehrensis var. hassiaca (Krasske) Lange-Bertalot Johansen et al. 2004 

Chaetoceras amanita A. Cleve Kaczmarska & Rushforth 1983 

Chaetoceras elmorei Boyer Boyer 1927a 

Chaetoceros hohnii Graebner & Wujek Stoermer et al. 1999 

Chaetoceros muelleri Lemmermann Kociolek & Herbst 1992 

Chaetoceros muelleri var. subsalsum (Lemmermann) Johansen & Rushforth Rushforth & Johansen 1986 

Cocconeis americana Ehrenberg 1856 

Cocconeis amygdalina (Brebisson) Grunow Patrick & Reimer 1966 

Cocconeis borealis Ehrenberg 1 856 

Cocconeis decussata Ehrenberg 1856 

Cocconeis delalineata Hohn Patrick & Reimer 1966 

Cocconeis delapunctata Hohn Patrick & Reimer 1966 

Cocconeis diminuta Pantocsek Stoermer & Kreis 1978 

Cocconeis diminuta var. aegagropilae Murobase Stoermer et al. 1999 

Cocconeis diruptoides Hustedt Stoermer & Kreis 1978 

Coccoensi distans Gregory Patrick & Reimer 1966 

Cocconeis disculus (Schumann) Cleve Stoermer & Kreis 1978 

Cocconeis disculus var. diminuta (Pantocsek) A. Cleve Stoermer & Kreis 1978 

Cocconeis elongata Ehrenberg 1856 

Cocconeis euglypta Ehrenberg Ehrenberg 1856 

Cocconeis finnica Ehrenberg Rushforth & Merkley 1988 

Cocconeis flexella (Kutzing) Cleve Stoermer & Kreis 1978 

Cocconeis fluviatilis Wallace Stoermer & Kreis 1978 

Cocconeis inusitatus Hohn Patrick & Reimer 1966 

Cocconeis japonica A. Schmidt Rushforth & Squires 1985 

Cocconeis klamathensis Sovereign Sovereign 1958 

Cocconeis limbata Ehrenberg Patrick & Reimer 1966 

Cocconeis lineata Ehrenberg Stoermer & Kreis 1978 

Cocconeis mexicana Ehrenberg Ehrenberg 1856 

Cocconeis mexicana Ehrenberg 1856 

Cocconeis mormonorum Ehrenberg Patrick & Reimer 1966 

Cocconeis neodiminuta Krammer Stoermer et al. 1999 

Cocconeis oblonga Kutzing Patrick & Reimer 1966 

Cocconeis patrickiae Reimer Patrick & Reimer 1966 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 41 1 



Name Publication 

Cocconeis pellucida Hantzsch Patrick & Reimer 1966 

Cocconeis pediculus Ehrenberg Stoermer & Kreis 1978 

Cocconeis pinnata Gregory Patrick & Reimer 1966 

Cocconeis placentula Ehrenberg Stoermer & Kreis 1978 

Cocconeis placentula var. euglypta (Ehrenberg) Cleve Stoermer & Kreis 1978 

Cocconeis placentula var. intermedia (Heribaud & Peragallo) Cleve Collins & Kalinsky 1977 

Cocconeis placentula var. klinoraphis Geitler Stoermer et al. 1999 

Cocconeis placentula var. hneata (Ehrenberg) Van Heurck Stoermer & Kreis 1978 

Cocconeis placentula var. rouxii (Heribaud & Brun) Cleve Stoermer et al. 1999 

Cocconeis praetexta Ehrenberg Patrick & Reimer 1966 

Cocconeis rouxii Heribaud & Brun Stoermer & Kreis 1978 

Cocconeis rhombea Ehrenberg Stoermer & Kreis 1978 

Cocconeis rugosa Sovereign Patrick & Reimer 1966 

Cocconeis scutellum Ehrenberg Stoermer & BCreis 1978 

Cocconeis scutellum var. japonica (A. Schmidt) Skvortzow Stoermer et al. 1999 

Cocconeis scutellum var. parva Grunow Stoermer & Kreis 1978 

Cocconeis scutellum f. par\'a Grunow in Van Heurck Patrick & Reimer 1966 

Cocconeis silicula (Ehrenberg) Cleve Kutkuhn 1958 

Cocconeis striata Ehrenberg 1856 

Cocconeis thumensis A. Mayer Stoermer & Kreis 1978 

Cocconeis thwaitesii W. Smith Stoermer & Kreis 1978 

Cocconeis transversalis Gregory Stoermer & Kreis 1978 

Cocconeis turgida Ehrenberg 1856 

Cocconeis undulata Ehrenberg Patrick & Reimer 1966 

Cocconema arcus Ehrenberg Patrick & Reimer 1975 

Cocconema asperum Ehrenberg 1856 

Cocconema australicum A. Schmidt Patrick & Reimer 1975 

Cocconema cistula Hemprich Stoermer & Kreis 1978 

Cocconema cistula Ehrenberg Cleve & Moller 

Cocconema comutum Ehrenberg Patrick & Reimer 1975 

Cocconema cymbiforme (Kutzing) Ehrenberg Rushforth & Squires 1985 

Cocconema eurypterum Ehrenberg 1856 

Cocconema excisum Kutzing H.L. Smith 1876-1888 (#81) 

978 
978 
975 
856 
856 
975 
978 
856 
856 
978 
978 
856 

978 
977 
977 



Cocconema fusidium Ehrenberg Stoermer & Kreis 

Cocconema gastroides (Kutzing) Pell Stoermer & Kreis 

Cocconema gibbum Ehrenberg Patrick & Reimer 

Cocconema gloeonema Ehrenberg 

Cocconema gracile Ehrenberg 

Cocconema helveticum (Kutzing) Grunow Patrick & Reimer 

Cocconema lanceolatum Ehrenberg Stoermer & Kreis 

Cocconema lunula Ehrenberg 

Cocconema mexicana Ehrenberg 

Cocconema parva W. Smith Stoermer & Kreis 

Cocconema scotica W. Wmith Stoermer & Kreis 

Cocconema subtile Ehrenberg 

Collectonema lacustre Van Heurck Stoermer & Kreis 

Collectonema minutum Collins & Kalinsky 

Collectonema vulgare Thwaites Collins & Kalinsky 

Coscinodiscus apiculatus Ehrenberg Rushforth & Merkley 

Coscinodiscus argus Ehrenberg Rushforth & Merkley 

Coscinodiscus asteromorphus Ehrenberg Stoermer & Kreis 

Coscinodiscus bodanica Schneider Prescott & Dillard 

Coscinodiscus catenata Brunnthaler Prescott & Dillard 

Coscinodiscus chambonis M. Peragallo & Heribaud Tempere & Peragallo 



978 
979 
979 
909 



4 1 2 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Name Publication 

Coscinodiscus curvatalus Grunow Stoermer & Kreis 1978 

Coscinodiscus decrescens Grunow Stoermer & Kreis 1978 

Coscinodiscus denarius A. Schmidt Czamecki & Blinn 1978 

Coscinodiscus lacustris Grunow Stoermer & Kreis 1978 

Coscinodiscus lanceolatum Ehrenberg Stoermer & Kreis 1978 

Coscinodiscus marginatus Ehrenberg Stoermer & Kreis 1978 

Coscinodiscus odontodiscus Grunow Rushforth & Merkley 1988 

Coscinodiscus pygmaeus var. micropunctatus M. Peragallo & Heribaud Drum 1981 

Coscinodiscus radiatus Ehrenberg Stoermer & Kreis 1978 

Coscinodiscus rothii (Ehrenberg) Grunow Stoermer & Kreis 1978 

Coscinodiscus rothii var. subsalsa (Juhl.-Dannf.) Hustedt Stoermer & Kreis 1978 

Coscinodiscus subsalsa Juhl.-Dannf Stoermer & Kreis 1978 

Coscinodiscus subtilis Ehrenberg Cambum 1982 

Coscinodiscus subtihs var. radiatus Hohn Hohn 1952 

Coscinodiscus tuberculatus Greville Stoermer & Kreis 1978 

Coscinodiscus woodwardii Tilden 1894-1909 (#367) 

Cosmioneis lundstroemii (Cleve in Cleve & Grunow) D.G Mann in Round et al Stoermer et al. 1999 

Cosmioneis pusilla (W. Smith) D.G Mann in Round et al Stoermer et al. 1999 

Craspedodiscus coscinodiscus Ehrenberg Stoermer et al. 1999 

Craspedodiscus microdiscus Ehrenberg Stoermer & Kreis 1978 

Craticuia accomoda (Hustedt) D.G. Mann in Round et al Stoermer et al. 1999 

Craticula acidochnata Lange-Bertalot & Metzeltin Siver et al. 2005 

Craticuia ambigua (Ehrenberg) D.G Mann in Round et al Stoermer et al. 1999 

Craticula citrus (Krassice) Reichardt Potapova & Charles 2003 

Craticuia cuspidata (Kiitzing) D.G Mann in Round et al Stoermer et al. 1999 

Craticula cuspidata var. major (Meister) Czamecki Stoermer et al. 1999 

Craticula halophihoides (Hustedt) Lange-Bertalot Siver et al. 2005 

Craticula perrotettii Grunow Lange-Bertalot 2001 

Craticula subhalophila (Hustedt) Lange-Bertalot Stoermer et al. 1999 

Craticula vixvisibilis (Hustedt) Lange-Bertalot Stoermer et al. 1999 

Ctenophora pulchella (Ralfs ex Kiitzing) Williams & Round Stoermer et al. 1999 

Ctenophora pulchella var. lacerata Stoermer et al. 1999 

Ctenophora pulchella var. lanceolata (O'Meara) Bukhtiyarova Stoermer et al. 1999 

Cyclostephanos costalimbus (Kob. & Kob.) Stoermer Hakansson & Theriot Stoermer et al. 1999 

Cyclostephanos delicatus (Genkal) Kling & Hakansson Hakansson & Kling 1990 

Cyclostephanos dubius (Fricke) Round in Theriot et al Stoermer et al. 1999 

Cyclostephanos invisitatus (Hohn & Hellerman) Theriot Stoermer & Hakansson Stoermer et al. 1999 

Cyclostephanos tholiformis Stoermer et al Hakansson & Kling 1990 

Cyclotella aliquantula Hohn & Hellerman Hohn & Hellerman 1963 

Cyclotella americana Fricke Stoermer et al. 1999 

Cyclotella antiqua W. Smith Stoermer & Kreis 1978 

Cyclotella arentii Kolbe Hamilton et al. 1992 

Cyclotella atomus Hustedt Stoermer & Kreis 1978 

Cyclotella berolinensis Ehrenberg Tilden 1894- 1909 (#367) 

Cyclotella bodanica Eulenstein Stoermer & Kreis 1978 

Cyclotella bodanica var. lemanensis O. Miiller Clark & Rushforth 1977 

Cyclotella bodanica var. michiganensis Skvortzow 1937 Stoermer & Kreis 1978 

Cyclotella bodanica var. stellata Skvortzow 1937 Stoermer & Kreis 1978 

Cyclotella caspia Grunow Stoermer & Kreis 1978 

Cyclotella catenata Brun Stoermer & Kreis 1978 

Cyclotella chaetoceras Lemmermann Stoermer & Kreis 1978 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 413 



Name Publication 

Cyclotella choctawhatcheeana Prasad Carvallo et al. 1995 

Cyclotella comensis Grunow Stoermer & Kreis 1978 

Cyclotella comta (Ehrenberg) Kiitzing Stoermer & Kreis 1978 

Cyclotella comla var. bodanica Grunow Stoermer & Kreis 1978 

Cyclotella comta var. glabriuscula Grunow Stoermer & Kreis 1978 

Cyclotella comta var. oligactis (Ehrenberg) Grunow Stoermer & Kreis 1978 

Cyclotella comta var. paucipunctata Grunow Stoermer & Kreis 1978 

Cyclotella comta var. radiosa Grunow Patrick 1945 

Cyclotella cryptica Reimann et al Stoermer & Kreis 1978 

Cyclotella cyclopunctata Hakansson & Carter Stoermer et al. 1999 

Cyclotella delicatula Hustedt Stoermer et al. 1999 

Cyclotella distinguenda Hustedt Stoermer et al. 1999 

Cyclotella distinguenda var. unipunctata (Hustedt) Hakansson Stoermer et al. 1999 

Cyclotella dubia Hilse Stoermer & Kreis 1978 

Cyclotella facetia Hohn & Hellerman Stoermer & Kreis 1978 

Cyclotella florida Voigt Patrick 1968 

Cyclotella gamma Sovereign Sovereign 1963 

Cyclotella glomerata Bachmann Stoermer & Kreis 1978 

Cyclotella krammeri Hakansson Stoermer et al. 1999 

Cyclotella kuetzingiana Thwaites Stoermer & Kreis 1978 

Cyclotella kuetzingiana var. planetophora Fricke Stoermer & Kreis 1978 

Cyclotella kuetzingiana var. radiosa Fricke Stoermer & Kreis 1978 

Cyclotella kuetzingiana var. schumannii Grunow Gaufin et al. 1976 

Cyclotella melosiroides (Kirchner) Lemmermann Stoermer & Kreis 1978 

Cyclotella meneghiniana Kiitzing Stoermer & Kreis 1978 

Cyclotella meneghiniana var. plana Fricke Stoermer & Kreis 1978 

Cyclotella meneghiniana var. pumila (Grunow ex Van Heurck) Hustedt Grimes & Rushforth 1982 

Cyclotella meneghiniana var. rectangulata Patrick 1945 

Cyclotella meneghiniana var. stelligera Cleve & Grunow in Cleve Boyer 1927a 

Cyclotella meneghiniana var. stellulifera Cleve & Grunow Stoermer & Kreis 1978 

Cyclotella michiganiana Skvortzow 1937 Stoermer & Kreis 1978 

Cyclotella minutula Amott H.L. Smithl876-1888 (#107) 

Cyclotella nana Hustedt Stoermer & Kreis 1978 

Cyclotella ocellata Pantocsek Stoermer & Kreis 1978 

Cyclotella operculata (Agardh) Kutzing Stoermer & Kreis 1978 

Cyclotella operculata W. Smith H.L. Smith 1876-1888 (#107) 

Cyclotella operculata var. unipunctata (Fricke) Hustedt Stoermer et al. 1999 

Cyclotella oregonica H.L. Smith H.L. Smith 1876-1888 (#504) 

Cyclotella perforata M. Peragallo Tempere & Peragallo 1909 

Cyclotella planktonica Brunnthaler Stoermer & Kreis 1978 

Cyclotella pseudostelligera Hustedt Stoermer & Kreis 1978 

Cyclotella pseudosteolligera f. parva Czamecki & Blinn Czamecki 1979 

Cyclotella quadriiunta (Schroeter) Hustedt Stoermer & Kreis 1978 

Cyclotella radiosa Grunow Stoermer et al. 1999 

Cyclotella rossii Hakansson Stoermer et al. 1999 

Cyclotella rotula Kutzing Stoermer & Kreis 1978 

Cyclotella seratula Hohn & Hellerman Hohn & Hellerman 1963 

Cyclotella socialis Schutt Stoermer et al. 1999 

Cyclotella spinosa Schumann Stoermer & Kreis 1978 

Cyclotella stelligera (Cleve & Grunow) Van Heurck Stoermer & Kreis 1978 

Cyclotella stelligera var. tenuis Hustedt Dodd 1987 

Cyclotella striata (Kutzing) Grunow Stoermer & Kreis 1978 

Cyclotella striata var. ambigua Grunow Collins & Kalinsky 1977 

Cyclotella striata var. bipunctata Fricke Stoermer & Kreis 1978 

Cyclotella striata var. mesoleia Tempere & Peragallo 1913 

Cyclotella temperei M. Peragallo & Heribaud Stoermer & Kreis 1978 

Cyclotella terryana Tempere & Peragallo Tempere & Peragallo 1909 



414 



PROCEEDINGS OF THE CALIFORNIA AC ADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 



Name 



Publication 



Cyclotella thienemannii var. minuscula Jurilj Stoermer et al. 1999 

Cyclotella wolterecki Hustedt Stoermer & Kreis 1978 

Cyclotubicoalistus undatus Stoermer et al Stoermer et al. 1990 

Cylindrotheca gracilis (Brebisson ex Kiitzing) Grunow Cambum 1982 

Cymatopleura angulata Greville Stoermer & Kreis 1978 

Cymatopleura apiculata W. Smith Stoermer & Kreis 1978 

Cymatopleura campylodiscus J.W. Bailey H.L. Smith 1876-1888 (#641) 

Cymatopleura cochlea Brun Stoermer & Kreis 1978 

Cymatopleura elliptica (Brebisson & Godey) W. Smith Stoermer & Kreis 1978 

Cymatopleura elliptica var. constricta Grunow Rushforth & Squires 1985 

Cymatopleura elliptica var. hibemica (W. Smith) Van Heurck Stoermer & Kreis 1978 

Cymatopleura elliptica var. nobilis (Hantzsch) Hustedt Dodd 1987 

Cymatopleura elliptica f. spiralis Boyer Stoermer & Kreis 1978 

Cymatopleura hibemica W. Smith Stoermer & Kreis 1978 

Cymatopleura hibemica var. rhombica H.H. Chase Stoermer & Kreis 1978 

Cymatopleura librile (Ehrenberg) Pantocsek Cambum 1982 

Cymatopleura mannii M. Peragallo in Tempere & Peragallo Tempere & Peragallo 1909 

Cymatopleura solea (Brebisson & Godey) W. Smith Stoermer & Kreis 1978 

Cymatopleura solea var. apiculata (W. Smith) Ralfs Stoermer & Kreis 1978 

Cymatopleura solea var. clavata O. Miiller Stoermer & Kreis 1978 

Cymatopleura solea var. gracilis Granow Stoermer et al. 1999 

Cymatopleura solea var. palffyi (Pantocsek) Cleve-Euler Rushforth & Squires 1985 

Cymatopleura solea var. pfuhlii Torka Stoermer & Kreis 1978 

Cymatopleura solea var. regula (Ehrenberg) Grunow Stoermer & Kreis 1978 

Cymatopleura solea var. subconstricta O. Muller Stoermer et al. 1999 

Cymatopleura solea var. vulgaris Meister Tiffany & Britton 1952 

Cymatopleura spiralis H.H. Chase Stoermer & Kreis 1978 

Cymbella acuta (A. Schmidt) Cleve Patrick & Reimer 1975 

Cymbella acutiuscula Cleve Stoermer & Kreis 1978 

Cymbella aequalis W. Smith Stoermer & Kreis 1978 

Cymbella aequalis var. subaequalis Gmnow Collins & Kalinsky 1977 

Cymbella affmis Kiitzing Stoermer & Kreis 1978 

Cymbella alpestris Krammer Krammer 2002 

Cymbella americana A.Schmidt Tempere & Peragallo 1908 

Cymbella americana f. minor Van Heurck & Gmnow 1882-1885 (#138) 

Cymbella amphicephala Nageli Stoermer & Kreis 1978 

Cymbella amphicephala var. subundulata Cleve Stoermer & Kreis 1978 

Cymbella amphio.xys (Kiitzing) Cleve Bateman & Rushforth 1984 

Cymbella anglica Lagerstedt Stoermer & Kreis 1978 

Cymbella angustata (W. Smith) Cleve Stoermer & Kreis 1978 

Cymbella aspera (Ehrenberg) H. Peragallo Stoermer & Kreis 1978 

Cymbella aspera var. minor (Van Heurck) Cleve Stoermer & Kreis 1978 

Cymbella australica (A. Schmidt) Conger Hohn 1951 

Cymbella austriaca Gmnow Stoermer & Kreis 1978 

Cymbella bonne\ illensis Setty Rushforth & Merkley 1988 

Cymbella borealis Cleve Cambum & Charles 2000 

Cymbella brehmii Hustedt Stoermer & Kreis 1978 

C\mbella buechleri Krammer Krammer 2002 

Cymbella caespitosa Bmn Stoermer & Kreis 1978 

Cymbella caespitosum Kiitzing Aubert 1 895 

C\mbella capitata M. Peragallo Patrick & Reimer 1975 

C\mbella cesatii (Rabenhorst) Gmnow Stoermer & Kreis 1978 

Cymbella cesati var. linearis Reimer Reimer 1961 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 415 



Name Publication 

Cymbella cistula (Ehrenherg) Kirchner Stoermer & Kreis 1978 

Cymbella cistula var. crassa Tempere & Peragallo Tempere & Peragallo 1909 

Cymbella cistula var. fusidium Tempere & Peragallo 1909 

Cymbella cistula var. gibbosa Brun Stoermer & Kreis 1978 

Cymbella cistula var. gracilis Hustedt Patrick & Reimer 1975 

Cymbella cistula var. maculata (Kutzing) Van Heurck Stoermer & Kreis 1978 

Cymbella cistula var. truncata Brun Stoermer & Kreis 1978 

Cymbella clausii Van Landingham Cambum 1982 

Cymbella couleensis Sovereign Sovereign 1963 

Cymbella cucumis var. delicata Tempere & Peragallo Tempere & Peragallo 1909 

Cymbella curta A. Schmidt Boyer 1927b 

Cymbella cuspidata Kiitzing Stoermer & Kreis 1978 

Cymbella cuspidata var. lanceolata May Clark & Rushforth 1977 

Cymbella cuspidata var. schulzii A. Cleve Stoermer & Kreis 1978 

Cymbella cuspidata f. impressa Fusey Stoermer et al. 1999 

Cymbella cymbiformis Agardh Stoermer & Kreis 1978 

Cymbella cymbiformis var. nonpunctata Fontell Collins & Kalinsky 1977 

Cymbella cymbiformis var. parra (W. Smith) Van Heurck Myers 1898b 

Cymbella delicatula Kutzing Stoermer & Kreis 1978 

Cymbella delicatula var. intermedia McCall Patrick & Reimer 1975 

Cymbella descripta (Hustedt) Krammer Cambum & Charles 2000 

Cymbella designata Krammer Stoermer et al. 1999 

Cymbella diluviana (Krasske) Florin Stoermer & Kreis 1978 

Cymbella dissimilis M. Peragallo in Tempere & Peragallo Tempere & Peragallo 1909 

Cymbella dorsirostrata Krammer Krammer 2002 

Cymbella duplopunctata Krammer Krammer 2002 

Cymbella ehrenbergii Kutzing Stoermer & Kreis 1978 

Cymbella ehrenbergii var. hungarica Pantocsek Clark & Rushforth 1977 

Cymbella ehrenbergii var. minor Tempere & Peragallo 1909 

Cymbella elizabethana Krammer Krammer 2002 

Cymbella elginsis Krammer Stoermer et al. 1999 

Cymbella excisa Kutzing Boyer 1927b 

Cymbella fluminea Patrick & Freese Patrick & Reimer 1975 

Cymbella fonticola Hstedt Collins & Kalinsky 1977 

Cymbella formosa Hustedt Hustedt 1955 

Cymbella gastroides Kutzing Stoermer & Kreis 1978 

Cymbella gasteroides (Kiitzing) Kutzing Rushforth & Merkley 1988 

Cymbella gerloffii Van Landingham Camburn 1982 

Cymbella gibba J.W. Bailey Patrick & Reimer 1975 

Cymbella gibberula Hustedt Krammer 2002 

Cymbella gracilis (Rabenhorst) Cleve Stoermer & Kreis 1978 

Cymbella gracilis var. lunata W. Smith Whitford & Schumacher 1973 

Cymbella hauckii Van Heurck Stoermer & Kreis 1978 

Cymbella hebridica Grunow ex Cleve Stoermer & Kreis 1978 

Cymbella helvetica Kutzing Stoermer & Kreis 1978 

Cymbella heteropleura (Ehrenberg) Kutzing Boyer 1927b 

Cymbella heteropleura var. minor Cleve Clark & Rushforth 1977 

Cymbella heteropleura var. subrostrata Cleve Patrick & Reimer 1975 

Cymbella heteropleura var. symmetrica Boyer Boyer 1927b 

Cymbella hohnii Van Landingham Camburn 1982 

Cymbella hungarica var. grunowii A. Cleve Clark & Rushforth 1977 

Cymbella hustedtii Krasske Stoermer & Kreis 1978 

Cymbella hybrida Grunow Stoermer & Kreis 1978 

Cymbella hybridiformis Hustedt Patrick & Reimer 1975 

Cymbella inaequalis (Ehrenbergj Rabenhorst Stoermer & Kreis 1978 

Cymbella incerta (Grunow) Cleve Stoermer & Kreis 1978 

Cymbella incerta var. naviculacea Grunow Johansen et al. 1983 



416 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

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Name Publication 

Cymbella inelegans Cleve Patrick & Reimer 1975 

Cymbella janischii A. Schmidt Boyer 1927b 

Cymbella javanica Hustedt Cambum 1982 

Cymbella jordani Grunow Stoermer & Kreis 1978 

Cymbella kappii Cholnoky Dodd 1987 

Cymbella lacustris (Agardh) Cleve Boyer 1927b 

Cymbella laevis Nageli Stoermer & Kreis 1978 

Cymbella lanceolata (Agardh) Agardh Stoermer & Kreis 1978 

Cymbella lanceolata var. comuta (Ehrenberg) Grunow Krammer 2002 

Cymbella lanceolatum Ehrenberg Aubert 1 895 

Cymbella langii MacLaughlin & Andrews Krammer 2002 

Cymbella lata Grunow Stoermer & Kreis 1978 

Cymbella latens Krasske Stoermer & Kreis 1978 

Cymbella laubyi M. Peragallo & Heribaud Tempere & Peragallo 1912 

Cymbella leptoceros (Ehrenberg) Rabenhorst Stoermer & Kreis 1978 

Cymbella leptoceros var. angusta Grunow Boyer 1927b 

Cymbella leptoceros var. rostrata Hustedt Stoermer & Kreis 1978 

Cymbella lunata W. Smith Cambum 1982 

Cymbella maculata Kiitzing Stoermer & Kreis 1978 

Cymbella mexicana (Ehrenberg) Cleve Stoermer & Kreis 1978 

Cymbella mexicana var. janischii (A. Schmidt) Reimer Patrick & Reimer 1975 

Cymbella mexicana var. punctifera Krammer Krammer 2002 

Cymbella microcephala Grunow Stoermer & Kreis 1978 

Cymbella microcephala var. crassa Reimer Stoermer et al. 1999 

Cymbella minuta Hilse Stoermer & Kreis 1978 

Cymbella minuta var. pseudogracilis (Cholnoky) Reimer Stoermer & Kreis 1978 

Cymbella minuta var. silesiaca (Bleisch) Reimer Stoermer & Kreis 1978 

Cymbella minuta f. latens (Krasske) Reimer Stoermer & Kreis 1978 

Cymbella moelleriana Grunow Stoermer & Kreis 1978 

Cymbella muelleri Hustedt Stoermer & Kreis 1978 

Cymbella muelleri f. ventricosa (Tempere & Peragallo) Reimer Stoermer & Kreis 1978 

Cymbella naviculiformis Auerswald Stoermer & Kreis 1978 

Cymbella norvegica Grunow Stoermer & Kreis 1978 

Cymbella norvegica f. minor Fusey Patrick & Reimer 1975 

Cymbella obtusa Gregory Stoermer & Kreis 1978 

Cymbella obtusa f. krasskei Foged Reimer 1961 

Cymbella obtusiuscula Kiitzing Stoermer & Kreis 1978 

Cymbella omata Hustedt Hustedt 1 93 1 

Cymbella para (W. Smith) Wolle Benson & Rushforth 1975 

Cymbella parva (W. Smith) Cleve Stoermer & Kreis 1978 

Cymbella parvula Krasske Stoermer & Kreis 1978 

Cymbella pediculus Kiitzing Stoermer & Kreis 1978 

Cymbella perfossilis Krammer Krammer 2002 

Cymbella perpusilla A. Cleve Cambum 1982 

Cymbella philadelphica Boyer Boyer 1927b 

Cymbella procera Hustedt Hustedt 1931 

Cymbella producta M. Peragallo Tempere & Peragallo 1908 

Cymbella prostrata (Berkeley) Cleve Stoermer & Kreis 1978 

Cymbella prostrata var. auerswaldii (Rabenhorst) Reimer Stoermer & Kreis 1978 

Cymbella protracta 0stmp Rushforth & Squires 1985 

Cymbella proxima Reimer Stoermer & Kreis 1978 

Cymbella proxima f. gravida Reimer Patrick & Reimer 1975 

Cymbella pusilla Gmnow Stoermer & Kreis 1978 

Cymbella rabenhorstii R. Ross Stoermer et al. 1999 

Cymbella rainierensis Sovereign Patrick & Reimer 1975 

Cymbella reinhardtii Gmnow Patrick & Reimer 1975 

Cymbella rhomboidea Boyer Stoermer & Kreis 1978 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 417 



Name Publication 

Cymbella robertii Krammer Krammer 2002 

Cymbella rotundata H.H. Chase Stoermer & Kreis 1978 

Cjinbella rugosa Hustedt Stoermer & Kreis 1978 

Cymbella rupicola Grunow Collins & Kalinsky 1977 

C>Tnbella rutmeri Hustedt Stoermer & Kreis 1978 

Cymbella ruttneri var. obtusa Hustedt Cambum 1982 

Cymbella schubartoides Cambum & Charles Camburn & Charles 2000 

Cymbella schmidtii Grunow Collins & Kalinsky 1977 

Cymbella scotica W. Smith Patrick 1945 

Cymbella silesiaca Bleisch Yearsley et al. 1992 

Cymbella similis Krasske Stoermer & Kreis 1978 

Cymbella sinuata Gregory Stoermer & Kreis 1978 

Cymbella sinuata var. antiqua (Grunow) Cleve Stoermer & Kreis 1978 

Cymbella sinuata var. ovata Hustedt Stoermer & Kreis 1978 

Cymbella sinuata f. antiqua (Grunow) Reimer Stoermer et al. 1999 

Cymbella sinuata f. ovata (Hustedt) Hustedt Stoermer & Kreis 1978 

Cymbella stauroneiformis Lagestedt Boyer 1927b 

Cymbella stodderi Cleve Boyer 1927b 

Cymbella stomatophora Grunow Stoermer & Kreis 1978 

Cymbella subaequalis Grunow Stoermer & Kreis 1978 

Cymbella subaequalis f. krasskei (Foged) Reimer Stoermer et al. 1999 

Cymbella subaspera var. salina Krammer Krammer 2002 

Cymbella subcistula Krammer Siver et al. 2005 

Cymbella subventricosa Cholnoky Stoermer & Kreis 1978 

Cymbella thumensis (A. Mayer) Hustedt Patrick & Reimer 1975 

Cymbella triangulum (Ehrenberg) Cleve Stoermer & Kreis 1978 

Cymbella tumida (Brebisson) Van Heurck Stoermer & Kreis 1978 

Cymbella tumida var. borealis (Grunow) Cleve Patrick & Reimer 1975 

Cymbella tumidula Grunow Stoermer & Kreis 1978 

Cymbella turgida Gregory Stoermer & Kreis 1978 

Cymbella turgida var. pseudogracilis Cholnoky Stoermer & Kreis 1978 

Cymbella mrgidula Grunow Stoermer & Kreis 1978 

Cymbella ventricosa Agardh Stoermer & Kreis 1978 

Cymbella ventricosa var. auerswaldii Meister Hohn & Hellerman 1963 

Cymbella ventricosa var. girodi (Heribaud) H. Kobayashi Patrick & Reimer 1975 

Cymbella ventricosa var. ovata f. minor Cleve-Euler Patrick & Reimer 1975 

Cymbella ventricosa var. silesiaca (Bleisch) Cleve Prescott & Dillard 1979 

Cymbellonitzschia diluviana Hustedt Stoermer & Kreis 1978 

Cymbopleura acutiuscula (Cleve) Krammer Krammer 2003 

Cymbopleura lata var. americana Krammer Krammer 2003 

Cymbopleura oregonica (Cleve) Krammer Krammer 2003 

Cymbopleura oregonica var. lata Krammer Krammer 2003 

Cymbopleura omata (Hustedt) Krammer Krammer 2003 

Cymbopleura peroregonica Krammer Krammer 2003 

Cymbopleura procera (Hustedt) Krammer Krammer 2003 

Cymbopleura subrostrata (Cleve) Krammer Krammer 2003 

Cystopleura argus fKutzing) Kuntze Elmore 1922 

Cystopleura gibba (Ehrenberg) Kuntze Stoermer & Kreis 1978 

Cystopleura gibberula fKiitzing) Kuntze Elmore 1922 

Cystopleura musculus (Kiitzing) Kuntze Elmore 1922 

Cystopleura musculus var. constricta (Brebisson) Van Heurck 

Cystopleura ocellata CEhrenberg) Kuntze Elmore 1922 

Cystopleura sorex (Kutzingj Kuntze Tilden 1894-1909 (#99) 

Cystopleura turgida CEhrenbergj Kuntze Stoermer & Kreis 1978 



4 1 8 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Name Publication 

Cystopleura ventricosa (Kiitzing) Elmore Elmore 1922 

Cystopleura zebra (Ehrenberg) Kuntze Elmore 1922 

Cystopleura zebra var. proboscidea (Kutzing) De Toni Patrick 1945 

Cystopleura zebra var. saxonica (Kutzing) De Toni Patrick 1945 

Decussata placenta (Ehrenberg) Lange-Bertalot in Metzeltin & Lange-Bertalot Johansen et al. 2004 

Delicata gerloffii (Van Landingham) Krammer Krammer 2003 

Denticula elegans Kutzing Stoermer & Kreis 1978 

Denticula elegans var. kittoniana (Grunow) DeToni Patrick & Reimer 1975 

Denticula elegans f. valida Pedicino Patrick & Reimer 1975 

Denticula frigida Kiitzing Stoermer & Kreis 1978 

Denticula kutzingii Grunow Cambum & Charles 2000 

Denticula lauta J.W. Bailey Stoermer & Kreis 1978 

Denticula palea Tilden 1894-1909 (#253) 

Denticula rainierensis Sovereign Patrick & Reimer 1975 

Denticula splendens Patrick & Reimer 1975 

Denticula subtilis Grunow Rushforth & Squires 1985 

Denticula tenuis Kutzing Stoermer & Kreis 1978 

Denticula tenuis var. crassula (Nageli) W. & GS. West Stoermer & Kreis 1978 

Denticula tenuis f. diminuta Manguin Dodd 1987 

Denticula tenuis var. frigida (Kutzing) Grunow Stoermer & Kreis 1978 

Denticula thermalis Kutzing Stoermer & Kreis 1978 

Denticula valida (Pedicino) Grunow in Van Heurck Tempere & Peragallo 1913 

Desmogonium guianense Ehrenberg Patrick 1946 

Desmogonium rabenhorstianum var. elongatum Patrick Whitford & Schumacher 1973 

Diadesmis confervacea Kutzing Stoermer et al. 1999 

Diadesmis contenta (Grunow ex Van Heurck) D.G Mann in Round et al Stoermer et al. 1999 

Diadesmis contenta var. biceps (Grunow in Van Heurck) Hamilton in Hamilton et al Hamilton et al. 1992 

Diadesmis gallica var. nitzschioides Grunow Cleve & Moller 1879 

Diadesmis peregrina W. Smith Stoermer et al. 1999 

Diadesmis perpusilla (Grunow) D.G Mann in Round et al Stoermer et al. 1999 

Diatoma anceps (Ehrenberg) Kirchner Stoermer & Kreis 1978 

Diatoma anceps var. capitatum Peragallo in Terry Boyer 1927a 

Diatoma anceps var. constricta Tempere & Peragallo Tempere & Peragallo 1912 

Diatoma anceps var. linearis M. Peragallo Stoermer & Kreis 1978 

Diatoma anceps var. mesodon (Ehrenberg) Grunow in Van Heurck Hamilton et al. 1992 

Diatoma ehrenbergii Kutzing Stoermer & Kreis 1978 

Diatoma elongata .^gardh Stoermer & Kreis 1978 

Diatoma elongatum (Lyngbye) Agardh Stoermer & Kreis 1978 

Diatoma elongatum var. minor Grunow Stoermer & Kreis 1978 

Diatoma elongatum van tenue (Agardh) Van Heurck Stoermer & Kreis 1978 

Diatoma elongatum \ar. tenuis (Agardh) Van Heurck Stoermer & Kreis 1978 

Diatoma hiemale (Roth) Heiberg Stoermer & Kreis 1978 

Diatoma hiemale var. mesodon (Ehrenberg) Grunow Stoermer & Kreis 1978 

Diatoma hyemale f. curta Patrick & Reimer 1966 

Diatoma mesodon (Ehrenberg) Kutzing Hamilton et al. 1992 

Diatoma moniliformis Kutzing Potapova & Charles 2002 

Diatoma stellaris Patrick & Reimer 1966 

Diatoma stellata Patrick & Reimer 1966 

Diatoma tenue Agardh Stoermer & Kreis 1978 

Diatoma tenue var. elongatum Lyngbye Stoermer & Kreis 1978 

Diatoma tenue var. pachycephala Grunow Stoermer & Kreis 1978 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 419 



Name Publication 



Diatoma vulgare Bor\ Stoermer & Kreis 

Diatoma vulgare var. breve Gninow Patrick & Reimer 

Diatoma vulgare var. capitulatum Grunow in Van Heurck Patrick & Reimer 

Diatoma vulgare var. ehrenhergii (Kiitzing) Grunow Stoermer & Kreis 

Diatoma vulgare var. grande (W. Smith) Grunow Stoermer & Kreis 

Diatoma vulgare var. ovalis (Fricke) Hustedt Stoermer & Kreis 

Diatoma vulgare var. pachycephala Grunow Gaufm 

Diatoma vulgare var. producta Grunow Stoermer & Kreis 

Diatoma vulgare var. linearis Van Heurck Stoermer & Kreis 

Diatomella balfouriana Greville Cambum 

Didymosphenia geminata (Lyngbye) M. Schmidt Stoermer & Kreis 

Diploneis boldtiana Cleve Stoermer & Kreis 

Diploneis domblittensis Grunow Stoermer & Kreis 

Diploneis elliptica (Kutzing) Cleve Stoermer & Kreis 

Diploneis elliptica var. ladogensis Cleve Stoermer et al. 

Diploneis elliptica var. pygmaea A. Cleve Stoermer & Kreis 

Diploneis fmnica (Ehrenberg) Cleve Stoermer & Kreis 

Diploneis fusca (Gregory) Cleve Johansen et al. 

Diploneis fusca var. delicata (A. Schmidt) Cleve Patrick & Reimer 

Diploneis interrupta (Kutzing) Cleve Stoermer & Kreis 

Diploneis marginestriata Hustedt Stoermer & Kreis 

Diploneis oblongella (Nageli) Ross Stoermer & Kreis 

Diploneis oblongella var. genuina Nageli Stoermer & Kreis 

Diploneis ocellata 0strup Stoermer et al. 

Diploneis oculata (Brebisson) Cleve Stoermer & Kreis 

Diploneis oculata var. linearis Gallik Prescott & Dillard 

Diploneis ostracodarum (Pantocsek) Jurilj Patrick & Reimer 

Diploneis ovalis (Hilse) Cleve Stoermer & Kreis 

Diploneis ovalis var. oblongella (Naegeli) Cleve Hohn 

Diploneis papula (A. Schmidt) Cleve Stoermer & Kreis 

Diploneis parma Cleve Stoermer & Kreis 

Diploneis peterseni Hustedt Stoermer & Kreis 

Diploneis pseudovalis Hustedt Stoermer & Kreis 

Diploneis puella (Schumann) Cleve Stoermer & Kreis 

Diploneis smithii (Brebisson) Cleve Stoermer & Kreis 

Diploneis smithii var. dilatata (M. Peragallo) Boyer Cambum 

Diploneis smithii var. pumila (Grunow) Hustedt Stoermer & Kreis 

Diploneis smithii f. rhombica Mereschkowsky Kaczmarska & Rushforth 

Diploneis subovalis Cleve Stoermer & Kreis 



978 
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966 
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978 
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978 
979 
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978 
983 
978 



Distrionella incognita (Reichardt) Williams Morales et al. 2005 

Ellerbeckia arenaria (Moore ex Ralfs) R.M. Crawford Stoermer et al. 1999 

Encyonema auerswaldii Rabenhorst Stoermer et al. 1999 

Encyonema brehmii (Hustedt) D.G. Mann in Round et al Stoermer et al. 1999 

Encyonema caespitosum Kiitzing Stoermer & Kreis 1978 

Encyonema evergladianum Krammer Krammer 1997b 

Encyonema formosum (Hustedt) D.G. Mann Krammer 1997a 

Encyonema gaeumannii (Meister) Krammer Siver et al. 2005 

Encyonema gibbum (J.W. Bailey) Krammer Krammer 1997a 

Encyonema gracile Rabenhorst Stoermer & Kreis 1978 

Encyonema hebridicum Grunow ex Cleve Krammer 1997b 

Encyonema hohnii (Van Landingham) Krammer Krammer 1997b 



420 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Name Publication 

Encyonema inelegans (Cleve) Mills Krammer 1997a 

Encyonema lacustre (C. Agardh) D.G. Mann in Round et al Stoermer et al. 1999 

Encyonema latens (Krasske) D.G Mann in Round et al Stoermer et al. 1999 

Encyonema lunatum (W. Smith) Van Heurck Krammer 1997a 

Encyonema minutum (Hilse ex Rabenhorst) D.G. Mann in Round et al Stoermer et al. 1999 

Encyonema minutum var. pseudogracilis (Cholnoky) Czamecki Stoermer et al. 1999 

Encyonema muelleri (Hustedt) D.G Mann in Round et al Stoermer et al. 1999 

Encyonema muelleri f. ventricosa (Tempere & M. Peragallo) Czamecki Stoermer et al. 1999 

Encyonema neomesianum Krammer Johansen et al. 2004 

Encyonema norvegicum (Grunow) Mills Krammer 1997b 

Encyonema parallelum M. Peragallo in Tempere & Peragallo Tempere & Peragallo 1909 

Encyonema perpusillum (A. Cleve) Mann in Round et al Hamilton et al. 1992 

Encyonema prostratum Ralfs Stoermer & Kreis 1978 

Encyonema rugosum (Hustedt) D.G Mann Krammer 1997a 

Encyonema silesiacum (Bleisch ex Rabenhorst) D.G Mann in Round et al Stoermer et al. 1999 

Encyonema silesiacum var. elegans Krammer Krammer 1997a 

Encyonema temperei Krammer Krammer 1997b 

Encyonema thermale Krammer Krammer 1997b 

Encyonema triangulatum Ktitzing Stoermer et al. 1999 

Encyonema trianguliforme Krammer Krammer 1997a 

Encyonema triangulum Kutzing Stoermer & Kreis 1978 

Encyonema turgidum (Gregory) Grunow Stoermer & Kreis 1978 

Encyonema turgidum var. hebridicum Cleve & Moller 1879 

Enyconema turgidum var. ventricosa Tempere & Peragallo Tempere & Peragallo 1889-1895 

Encyonema ventricosa Ktitzing Stoermer & Kreis 1978 

Encyonema ventricosum var. angusta Krammer Siver et al. 2005 

Encyonema ventricosum var. stricta Tempere & Peragallo 1909 

Encyonema yellowstonianum Krammer Krammer 1997a 

Encyonopsis cesatii (Rabenhorst) Krammer Krammer 1997b 

Encyonposis floridana Krammer Krammer 1997b 

Encyonopsis kriegeri var. fossilis Krammer Krammer 1997b 

Encyonopsis microcephala (Grunow) Krammer Krammer 1997b 

Encyonopsis radialis Krammer Krammer 1997b 

Encyonopsis stodderi (Cleve) Krammer Krammer 1997b 

Encyonopsis subminuta Krammer in Reichardt & Krammer Siver et al. 2005 

Encyonopsis subspicula Krammer Krammer 1997b 

Encyonopsis substodderi Krammer Krammer 1997b 

Encyonopsis symmetrica Krammer Krammer 1997b 

Entomoneis alata (Ehrenberg) Ehrenberg Stoermer et al. 1999 

Entomoneis omata (J.W. Bailey) Reimer Stoermer & Kreis 1978 

Entomoneis paludosa (W. Smith) Reimer Cambum 1982 

Entomoneis paludosa var. duplex (Donkin) Cleve Kaczmarska & Rushforth 1983 

Entomoneis pulchra (J.W. Bailey) Reimer Patrick & Reimer 1975 

Entomoneis robusta (McCall) Reimer Patrick & Reimer 1975 

Epithemia adnata (Ktitzing) Brebisson Collins & Kalinsky 1977 

Epithemia adnata var. minor (M. Peragallo & Heribaud) Patrick Stoermer et al. 1999 

Epithemia adnata var. porcellus (Kutzing) Patrick Collins & Kalinsky 1977 

Epithemia adnata var. proboscidea (Kutzing) Patrick Stoermer et al. 1999 

Epithemia adnata var. saxonica (Ktitzing) Patrick Stoermer et al. 1999 

Epithemia alpestris W. Smith Stoermer & Kreis 1978 

Epithemia amphicephala Grunow Tempere & Peragallo 1909 

Epithemia andrewsii Stoermer & Yang Stoermer & Kreis 1978 

Epithemia argus (Ehrenberg) Ktitzing Stoermer & Kreis 1978 

Epithemia argus var. alpestris Grunow Stoermer & Kreis 1978 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 421 



Name Publication 

Epithemia argus var. amphicephala Gmnow Stoermer & Kreis 1978 

Epithemia argus var. longicomis (Ehrenberg) Grunow Stoermer & Kreis 1978 

Epithemia argus var. protracta A. Mayer Patrick & Reimer 1975 

Epithemia emarginata Andrews Stoermer & Kreis 1978 

Epithemia frickei Krammer Stoermer et al. 1999 

Epithemia gibba Kiitzing Stoermer & Kreis 1978 

Epithemia gibba var. parallela Grunow Stoermer et al. 1999 

Epithemia gibba var. ventricosa (Kutzing) Grunow Stoermer & Kreis 1978 

Epithemia gibberula (Ehrenberg) Kutzing Stoermer & Kreis 1978 

Epithemia gibberula var. producta Grunow Boyer 1927b 

Epithemia gibberula var. protracta Tempere & Peragallo 1913 

Epithemia hyndmanii W. Smith Stoermer & Kreis 1978 

Epithemia hyndmanii var. capitata M. Peragallo Patrick & Reimer 1975 

Epithemia hyndmanii var. perlonga Pantocsek Tempere & Peragallo 1908 

Epithemia intermedia Fricke Stoermer & Kreis 1978 

Epithemia muelleri Fricke Stoermer & Kreis 1978 

Epithemia ocellata (Ehrenberg) Kutzing Stoermer & Kreis 1978 

Epithemia reicheltii Fricke Stoermer & Kreis 1978 

Epithemia rupestris W. Smith H.L. Smith 1876-1888 (#152) 

Epithemia smithii Camithers Stoermer & Kreis 1978 

Epithemia sorex Kutzing Stoermer & Kreis 1978 

Epithemia truncata M. Peragallo Patrick & Reimer 1975 

Epithemia truncata var. debilis M. Peragallo Patrick & Reimer 1975 

Epithemia mrgida (Ehrenberg) Kutzing Stoermer & Kreis 1978 

Epithemia mrgida var. granulata (Ehrenberg) Brun Stoermer & Kreis 1978 

Epithemia turgida var. plicata Meister 

Epithemia turgida var. vertagus Tempere & Peragallo 1 909 

Epithemia turgida var. westermannii (Ehrenberg) Grunow Stoermer & Kreis 1978 

Epithemia turgida var. zebrina (Ehrenberg) Rabenhorst Stoermer et al. 1999 

Epithemia ventricosa Ehrenberg Stoermer & Kreis 1978 

Epithemia westermannii var. stricta Tempere & Peragallo Tempere & Peragallo 1909 

Epithemia zebra (Ehrenberg) Kutzing Stoermer & Kreis 1978 

Epithemia zebra f. minor Tempere & Peragallo 1909 

Epithemia zebra var. porcellus (Kutzing) Grunow Stoermer & Kreis 1978 

Epithemia zebra var. proboscida Grunow Tempere & Peragallo 1908 

Epithemia zebra var. saxonica (Kiitzing) Grunow Stoermer & Kreis 1978 

Eucocconeis depressa (Cleve) Hustedt Stoermer & Kreis 1978 

Eucocconeis diluviana (Hustedt) Lange-Bertalot Siver et al. 2005 

Eucocconeis flexella (Kutzing) Hustedt Stoermer & Kreis 1978 

Eucocconeis flexella var. alpestris (Brun) Hustedt Stoermer & Kreis 1978 

Eucocconeis lapponica Hustedt Stoermer & Kreis 1978 

Eucocconeis lapponica var. ninckei (Guermeur & Manguin) Edlund Stoermer & Kreis 1978 

Eucocconeis minuta (Cleve) Cleve Stoermer & Kreis 1978 

Eunotia aduncus Hohn & Hellerman Hohn & Hellerman 1963 

Eunotia amphioxys Ehrenberg Patrick & Reimer 1966 

Eunotia arcubus Norpel-Schempp & Lange-Bertalot Stoermer et al. 1999 

Eunotia arcus Ehrenberg Stoermer & Kreis 1978 

Eunotia arcus var. bidens Grunow Stoermer & Kreis 1978 

Eunotia arcus var. curta (Grunow) Schonfeldt Patrick 1945 

Eunotia arcus var. fallax Hustedt Stoermer & Kreis 1978 

Eunotia arcus var. plicata (Brun) Heribaud Patrick & Reimer 1966 

Eunotia arcus var. minor Grunow Boyer 1927a 

Eunotia arcus var. tenella Grunow Tempere & Peragallo 1908 

Eunotia arcus var. uncinata (Ehrenberg) Grunow Patrick & Reimer 1966 

Eunotia argus Ehrenberg 1 856 



422 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Name Publication 

Eunotia bactriana Ehrenberg Boyer 1927a 

Eunotia batavica f. gamma Berg Patrick & Reimer 1966 

Eunotia biceps Ehrenberg Boyer 1927a 

Eunotia bidens Ehrenberg Stoermer et al. 1999 

Eunotia bidentula W. Smith Boyer 1927a 

Eunotia bidentata W. Smith Cleve & Moller 1879 

Eunotia bigibba Kiitzing Lawson & Rushforth 1975 

Eunotia bigibba var. pumila Grunow Cambum 1982 

Eunotia bihi Lowe & Kociolek Lowe & Kociolek 1984 

Eunotia bilunaris var. mucophila Lange-Bertalot & Norpel Dute et al. 2000 

Eunotia camelus Ehrenberg Boyer 1927a 

Eunotia camelus f. dentata Berg Patrick & Reimer 1966 

Eunotia Carolina Patrick Patrick 1958 

Eunotia catillifera Morrow in Morrow, Deason & Clayton Morrow et al. 1981 

Eunotia cistula Ehrenberg 1856 

Eunotia clavata Hustedt Hustedt 1913 

Eunotia clevei Grunow Patrick & Reimer 1966 

Eunotia collinsii KaUnsky Kalinsky 1984 

Eunotia compacta Hustedt Hustedt 1913 

Eunotia cordillera Hohn & Hellerman Hohn & Hellerman 1963 

Eunotia cristagalli Cleve Cambum 1982 

Eunotia curvata (Kiitzing) Lagerstedt Stoermer & Kreis 1978 

Eunotia curvata f. bergii Woodhead & Tweed Cambum & Charles 2000 

Eunotia curvata var. capitata Patrick Whitford & Schumacher 1973 

Eunotia curvata var. falcata (Brebisson) Berg Clark & Rushforth 1977 

Eunotia curvata var. subarcuata (Nageli) Woodhead & Tweed Cambum 1982 

Eunotia cygnus Ehrenberg Patrick & Reimer 1966 

Eunotia denticulata (Brebisson) Rabenhorst Cambum & Charles 2000 

Eunotia depressa Ehrenberg Patrick & Reimer 1966 

Eunotia didyma var. inflata Hustedt Hustedt 1913 

Eunotia dianae Ehrenberg 1856 

Eunotia diodon Ehrenberg Stoermer & Kreis 1978 

Eunotia diodon f. minor Tempere & Peragallo 1908 

Eunotia ehrenbergii Ralfs Boyer 1927a 

Eunotia edegans 0strup Whitford & Schumacher 1973 

Eunotia elegans 0strup Cambum 1982 

Eunotia elongata Rabenhorst Patrick & Reimer 1966 

Eunotia euryptera Ehrenberg 1856 

Eunotia exigua (Brebisson) Rabenhorst Stoermer & Kreis 1978 

Eunotia exigua var. bidens Hustedt Hamilton et al. 1992 

Eunotia exigua var. compacta Hustedt Cambum & Charles 2000 

Eunotia exigua var. tridentula 0strup Cambum & Charles 2000 

Eunotia exigua var. undulata Magdeburg Hamilton et al. 1992 

Eunotia faba Ehrenberg emend Van Heurck Patrick 1945 

Eunotia fallax A. Cleve Cambum 1982 

Eunotia fallax var. gracillima Krasske Cambum 1982 

Eunotia fallax var. groenlandica (Grunow) Lange-Bertalot & Norpel Cambum & Charles 2000 

Eunotia flectuosa (Brebisson) Gmnow Hohn & Hellerman 1963 

Eunotia flexuosa Brebisson Stoermer & Kreis 1978 

Eunotia tlexuosa var. eurj'cephala Gmnow Stoermer & Kreis 1978 

Eunotia formica Ehrenberg Stoermer & Kreis 1978 

Eunotia formica f. alpha Berg Patrick & Reimer 1966 

Eunotia formica f. beta Berg Patrick & Reimer 1966 

Eunotia gibba Ehrenberg Patrick & Reimer 1966 

Eunotia gibbemla Ehrenberg 1856 

Eunotia gibbosa Gmnow Boyer 1927a 

Eunotia glacialis Meister Hansmann 1973 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 423 



Name Publication 

Eunotia gracilis (Ehrenberg) Rabenhorst Stoermer & Kreis 1978 

Eunotia gracilis f. major (M. Peragallo) Heribaud Patrick & Reimer 1966 

Eunotia granulata Ehrenberg Stoermer & Kreis 1978 

Eunotia gratella f. beta Berg Patrick & Reimer 1966 

Eunotia hemicyclus (Ehrenberg) Ralfs Boyer 1927a 

Eunotia hendecaodon Ehrenberg 1856 

Eunotia hexaglyphis Ehrenberg Patrick 1958 

Eunotia hinziae Simonsen 1987 Simonsen 1987 

Eunotia hinziae var. diodon Simonsen 1987 Simonsen 1987 

Eunotia iatriaensis Foged Camburn & Charles 2000 

Eunotia impressa Ehrenberg Boyer 1927a 

Eunotia incisa W. Smith Stoermer & Kreis 1978 

Eunotia incur\ata Hustedt Hustedt 1913 

Eunotia indica Grunow Patrick 1945 

Eunotia inflata (Grunow) Norpel-Schempp & Lange-Bertalot Stoermer et al. 1999 

Eunotia intermedia (Krasske) Norpel-Schempp & Lange-Bertalot Stoermer et al. 1999 

Eunotia kentuccensis Ehrenberg 1856 



Eunot 
Eunoi 
Eunot 
Eunot 
Eunot 
Eunot 
Eunot 
Eunot 
Eunot 
Eunot 
Eunot: 
Eunot: 
Eunot 
Eunot 
Eunot 
Eunot: 
Eunot 
Eunot: 
Eunot: 
Eunot 
Eunot 
Eunot: 
Eunot: 
Eunot: 
Eunot 
Eunot 
Eunot 
Eunot 
Eunot 
Eunot 
Eunot 
Eunoti 
Eunoti 
Eunot: 
Eunot: 
Eunot: 
Eunot 
Eunot 
Eunot 
Eunot 



a kocheUensis O. Muller Patrick & Reimer 1966 

a lapponica Grunow ex A. Cleve Patrick & Reimer 1966 

a lata Hustedt Hustedt 1933b 

a librile Ehrenberg Patrick & Reimer 1966 

a longicomis Ehrenberg 1 856 

a luna Ehrenberg Boyer 1927a 

a luna var. intermedia Hustedt ex Simonsen 1987 Hustedt 1913 

a luna var. elongata Hustedt ex Simonsen 1987 Hustedt 1913 

a luna var. aequalis Hustedt ex Simonsen 1987 Hustedt 1913 

a luna var. globosa Hustedt ex Simonsen 1987 Hustedt 1913 

a luna var. trapezica Hustedt Hustedt 1913 

a lunaris (Ehrenberg) Grunow Stoermer & Kreis 1978 

a lunaris var. attenuata A. Berg Dixit & Smol 1995 

a lunaris var. capitata Hustedt Patrick 1945 

a lunaris var. excisa Grunow Stoermer & Kreis 1978 

a lunaris var. subarcuata (Naegeli) Grunow Hohn & Hellerman 1963 

a lunula Ehrenberg Patrick & Reimer 1966 

a major Rabenhorst Stoermer & Kreis 1978 

a major f. compacta Berg Patrick & Reimer 1966 

a major f. excelsa Berg Patrick & Reimer 1966 

a major var. impressa (W. Smith) Rabenhorst Stoermer & Kreis 1978 

a major f. plectrum Berg Patrick & Reimer 1966 

a major var. ventricosa A. Cleve Patrick & Reimer 1966 

a meisteri Hustedt Camburn 1982 

a meisteri var. bidens Hustedt Dixit & Smol 1995 

a microcephala Krasske Stoermer & Kreis 1978 

a microcephala var. tridentata (Mayer) Hustedt Camburn et al. 1978 

a minor Rabenhorst Tempere & Peragallo 1908 

a mira var. ovata A. Berg Bateman & Rushforth 1984 

a monodon Ehrenberg Stoermer & Kreis 1978 

a monodon f. curta Tempere & Peragallo 1909 

a monodon var. bidens (Gregory) Hustedt Scherer 1988 

a monodon var. constricta Cleve-Euler Whitford & Schumacher 1973 

a monodon var. major (W. Smith) Hustedt Stoermer & Kreis 1978 

a monodon var. major f. bidens W. Smith Hohn 195 1 

a monodontiforma Lange-Bertalot & Norpel Stoermer et al. 1999 

a mosis Ehrenberg Rushforth & Merkley 1988 

a naegelii Migula Stoermer & Kreis 1978 

a nivalis Hohn & Hellerman Hohn & Hellerman 1963 

a nodosa Ehrenberg Patrick & Reimer 1966 



424 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Name Publication 

Eunotia nymanniana Grunow Boyer 1927a 

Eunotia obesa var. wardii Patrick Patrick 1958 

Eunotia paludosa Grunow Potapova & Charles 2002 

Eunotia paludosa var. trinacria (Krasske) Norpel Cambum & Charles 2000 

Eunotia papilio Ehrenberg Hohn 195 1 

Eunotia paradoxa Ehrenberg Patrick & Reimer 1966 

Eunotia parallela Ehrenberg Stoermer & Kreis 1978 

Eunotia pectinalis (O. Muller) Rabenhorst Stoermer & Kreis 1978 

Eunotia pectinalis var. biarcuata Berg Whitford & Schumacher 1973 

Eunotia pectinalis f. curta Tempere & Peragallo 1909 

Eunotia pectinalis f. didymodon (Grunow) Berg Patrick & Reimer 1966 

Eunotia pectinalis var. elongatum Aubert 1895 

Eunotia pectinalis var. macilenta Grunow Van Heurck & Grunow 1882-1885 (#138) 

Eunotia pectinalis var. minor (Kutzing) Rabenhorst Stoermer & Kreis 1978 

Eunotia pectinalis var. minor f. impressa (Ehrenberg) Hustedt Patrick 1945 

Eunotia pectinahs var. recta Mayer ex Patrick Patrick 1945 

Eunotia pectinalis var. stricta Rabenhorst Boyer 1927a 

Eunotia pectinalis var. soleirolii (Kutzing) Boyer Boyer 1927a 

Eunotia pectinalis var. undulata (Ralfs) Rabenhorst Stoermer & Kreis 1978 

Eunotia pectinalis var. ventralis (Ehrenberg) Hustedt Dodd 1987 

Eunotia pectinalis var. ventricosa Grunow Stoermer & Kreis 1978 

Eunotia pectinalis f. minor (Dillwyn) Rabenhorst Stoermer & Kreis 1978 

Eunotia pentaglyphis Ehrenberg Boyer 1927a 

Eunotia perminuta (Grunow) Patrick Patrick 1958 

Eunotia perpusilla Grunow Stoermer & Kreis 1978 

Eunotia pirla Carter & Flower Carter & Flower 1988 

Eunotia pocosinensis Gaiser & Johansen Gaiser & Johansen 2000 

Eunotia polydentula var. perpusilla Patrick 1968 

Eunotia polyglyphis Ehrenberg Tempere & Peragallo 1913 

Eunotia praerupta Ehrenberg Stoermer & Kreis 1978 

Eunotia praerupta var. bidens (Ehrenberg) Grunow Stoermer & Kreis 1978 

Eunotia praerupta var. curta Grunow Patrick 1945 

Eunotia praerupta var. inflata Grunow Stoermer & Kreis 1978 

Eunotia praerupta var. laticeps f. curta Grunow Stoermer & Kreis 1978 

Eunotia praerupta var. monodon Aubert 1895 

Eunotia praerupta var. monodon f. polaris (A. Berg) Symoens Cambum & Charles 2000 

Eunotia praerupta-nana Berg Patrick & Reimer 1966 

Eunotia prionotus Ehrenberg Patrick & Reimer 1966 

Eunotia pseudolunaris Venkt Stoermer & Kreis 1978 

Eunotia pseudo-parallela f. alpha Berg Patrick & Reimer 1966 

Eunotia punctastriatum Cambum & Charles Cambum & Charles 2000 

Eunotia quartemaria Ehrenberg Patrick 1958 

Eunotia rabenhorstii var. monodon Gmnow Patrick & Reimer 1966 

Eunotia recta Hustedt 1913 Hustedt 1913 

Eunotia reicheltii Hustedt 1913 Hustedt 1913 

Eunotia reicheltii var. bidens Hustedt 1913 Hustedt 1913 

Eunotia reicheltii var. triodon Hustedt 1913 Hustedt 1913 

Eunotia rhomboidea Hustedt Cambum 1982 

Eunotia robusta Ralfs Boyer 1927a 

Eunotia robusta var. diadema Patrick & Reimer 1966 

Eunotia robusta var. heudecaodon Ralfs Aubert 1895 

Eunotia robusta var. tetraedron (Ehrenberg) Ralfs Stoermer & Kreis 1978 

Eunotia robusta var. triodon Ehrenberg Aubert 1895 

Eunotia rostellala Hustedt ex Patrick Patrick 1945 

Eunotia rostrata Ehrenberg 1 856 

Eunotia sarekensis var. pumila (Gmnow) S. Berg Clark & Rushforth 1977 

Eunotia schweikerdtii Cholnoky Cambum et al. 1978 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 425 



Name Publication 

Eunotia sepiena Ehrenberg Patrick 1945 

Eunotia septentrionalis 0strup Stoermer & Kreis 1978 

Eunotia serra Ehrenberg Stoermer & Kreis 1978 

Eunotia serra var. diadema (Ehrenberg) Pati-ick Cambum 1982 

Eunotia serraceniae Gaiser & Johansen Gaiser & Johansen 2000 

Eunotia sima Ehrenberg Hustedt 1913 

Eunotia soleirolii (Kiitzing) Rabenhorst Patrick 1958 

Eimotia sphaerula Ehrenberg Patrick & Reimer 1 966 

Eunotia st.antonii Ehrenberg 1 856 

Eunotia stevensonii Boyer Boyer 1927a 

Eunotia submonodon Hustedt 1913 Hustedt 1913 

Eunotia sudetica O. Miiller Cambum 1982 

Eunotia suecica A. Cleve Patrick 1945 

Eunotia tauntoniensis Hustedt Boyer 1927a 

Eunotia tenella (Grunow) Hustedt Stoermer & Kreis 1978 

Eunotia temaria Ehrenberg Patrick & Reimer 1966 

Eunotia testudinata Berg Kalinsky 1979 

Eunotia tetraodon Ehrenberg Boyer 1927a 

Eunotia tetraodon f. minuta Berg Patrick & Reimer 1966 

Eunotia torula Hohn Patrick & Reimer 1966 

Eunotia tridentula Ehrenberg Boyer 1927a 

Eunotia tridentula var. perminuta Grunow Hohn 195 1 

Eunotia triodon Ehrenberg Boyer 1927a 

Eunotia trinacria Krasske Stoermer & Kreis 1978 

Eunotia trinacria var. undulata Hustedt Cambum 1982 

Eunotia turgida Ehrenberg 1856 

Eunotia uncinata Ehrenberg Cleve & Moller 1879 

Eunotia undulata Ralfs Tempere & Peragallo 1913 

Eunotia valida Hustedt Stoermer & Kreis 1978 

Eunotia vanheurckii Patrick Stoermer & Kreis 1978 

Eunotia vanheurckii var. intermedia (Krasske) Patrick Stoermer & Kreis 1978 

Eunotia varioundulata Norpel & Lange-Bertalot in Lang-Bertalot et al Siver et al. 2005 

Eunotia veneris (Kutzing) De Toni Boyer 1927a 

Eunotia ventralis Ehrenberg Hohn 195 1 

Eunotia westermanni Ehrenberg 1856 

Eunotia woleirotii (Kutzing) Rabenhorst Stoermer & Kreis 1978 

Eunotia zasuminensis (Cabejszekowna) Komer Kalinsky 1984 

Eunotia zasuminensis var. minor Kalinsky Kalinsky 1984 

Eunotia zebra Ehrenberg 1 856 

Eunotia zebrina Ehrenberg Stoermer & Kreis 1978 

Eunotia zygodon Ehrenberg Boyer 1927a 

Eunotia zygodon var. elongata Hustedt Gaiser & Johansen 2000 

Fallacia fracta (Hustedt ex Simonsen) D.G Mann in Round et al Stoermer et al. 1999 

Fallacia helensis (Schulzj D.G Mann in Round et al Stoermer et al. 1999 

Fallacia indifferens (Hustedt) D.G. Mann in Round et al Johansen et al. 2004 

Fallacia insociabilis (Krasske) D.G. Mann in Round et al Stoermer et al. 1999 

Fallacia monoculata (Hustedt) D.G. Mann in Round et al Stoermer et al. 1999 

Fallacia omissa (Hustedt) D.G. Mann in Round et al Stoermer et al. 1999 

Fallacia pseudomuralis (Hustedt) D.G. Mann in Round et al Stoermer et al. 1999 

Fallacia pygmaea (Kutzing) D.G. Mann in Round et al Stoermer et al. 1999 

Fallacia subhamulata (Grunow in Van Heurck) D.G. Mann in Round et al Stoermer et al. 1999 

Fallacia submitis (Hustedt) D.G Mann in Round et al Stoermer et al. 1999 

Fallacia tenera (Hustedt) D.G. Mann Potapova & Charles 2002 

Fallacia vitrea (0strup) D.G. Mann in Round et al Johansen et al. 2004 

Fistulifera saprophila (Lange-Bertalot & Bonik) Lange-Bertalot Johansen et al. 2004 



426 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Name Publication 



Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi: 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Fragi 
Frasi 



aria acidobionta Charles Charles 1986 

aria acuta Ehrenberg Stoermer & Kreis 1978 

aria aequalis Heiberg Tempere & Peragallo 1909 

aria aequalis var. major Tempere & Peragallo Tempere & Peragallo 1909 

aria aequalis var. producta Lagerstedt Tempere & Peragallo 1908 

aria arcus (Ehrenberg) Cleve Stoermer & Kreis 1978 

aria atomus Hustedt Stoermer & Kreis 1978 

aria bicapitata A. Mayer Stoermer & Kreis 1978 

aria biceps Ehrenberg Patrick & Reimer 1966 

aria bipunctata Ehrenberg Stoermer & Kreis 1978 

aria bidens Heiberg Boyer 1927a 

aria bituminosa Pantocsek Tempere & Peragallo 1913 

aria brevistriata Grunow Stoermer & Kreis 1978 

aria brevistriata var. binodis (Pantocsek) A. Cleve Stoermer et al. 1999 

aria brevistriata var. capitata Heribaud Stoermer & Kreis 1978 

aria brevistriata var. inflata (Pantocsek) Hustedt Stoermer & Kreis 1978 

aria brevistriata var. inflata f. curta Skvortzow Reimer 1961 

aria capucina Desm Stoermer & Kreis 1978 

aria capucina var. acuta Grunow Boyer 1927a 

aria capucina var. acuminata Grunow Boyer 1927a 

aria capucina var. familiaris (Kutzing) Hamilton & Siver in Siver et al Siver et al. 2005 

aria capucina var. gracilis Morales 2001 

aria capucina var. lanceolata Grunow Stoermer & Kreis 1978 

aria capucina var. mesolepta Rabenhorst Stoermer & Kreis 1978 

aria capucina var. vaucheriae (Kutzing) Lange-Bertalot Kalinsky 1982 

aria constricta Ehrenberg Hansmann 1973 

aria constricta f. stricta (A. Cleve) Hustedt Stoermer & Kreis 1978 

aria constricta var. tetranodis (A. Cleve) Hustedt Hamilton et al. 1992 

aria constricta var. trinodis Hustedt Hamilton et al. 1992 

aria construens (Ehrenberg) Grunow Stoermer & Kreis 1978 

aria construens var. bigibba A. Cleve Stoermer & Kreis 1978 

aria construens var. binodis (Ehrenberg) Grunow Stoermer & Kreis 1978 

aria construens var. capitata Heribaud Stoermer & Kreis 1978 

aria construens var. genuina Tempere & Peragallo 1895 

aria construens var. exigua (W. Smith) Schulz Patrick & Reimer 1966 

aria construens var. javanica Hustedt Camburn et al. 1978 

aria construens var. minuta Tempere & Peragallo Stoermer & Kreis 1978 

aria construens var. pumila Grunow Stoermer & Kreis 1978 

aria construens var. subsalina Hustedt Stoermer & Kreis 1978 

aria construens var. venter (Ehrenberg) Grunow Stoermer & Kreis 1978 

aria construens var. venter f. pusilla Grunow Clark & Rushforth 1977 

aria crotonensis Kitton Stoermer & Kreis 1978 

aria crotonensis var. oregona Sovereign Stoermer & Kreis 1978 

aria crotonensis var. prolongata Grunow Stoermer & Kreis 1978 

aria cuneata Ehrenberg Patrick & Reimer 1966 

aria dibolos Hohn & Hellerman Hohn & Hellerman 1963 

aria diophthalama (Ehrenberg) Ehrenberg Patrick & Reimer 1966 

aria elliptica Schumann Patrick 1945 

aria elliptica f. minor Patrick & Reimer 1966 

aria entomon Ehrenberg Stoermer & Kreis 1978 

aria eugramma Ehrenberg Patrick & Reimer 1966 

aria exigua Grunow Camburn & Charles 2000 

aria exiguiformis Lange-Bertalot Potapova & Charles 2003 

aria floridana Hanna Hein 1981 

aria glebula Hohn & Hellerman Hohn & Hellerman 1963 

aria gnathostoma Hohn Patrick & Reimer 1966 

aria 2racillima A. Maver Stoermer & Kreis 1978 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 427 



Name Publication 

Fragilaria harrisonii Grunow Stoermer & Kreis 1978 

Fragilaria harrisonii var. dubia Grunow Stoermer & Kreis 1978 

Fragilaria harrisonii var. rhomboides Grunow Stoermer & Kreis 1978 

Fragilaria heideni 0strup Stoermer & Kreis 1978 

Fragilaria heideni var. istvanffyi (Pantocsek) Hustedt Stoermer & Kreis 1978 

Fragilaria hungarica Pantocsek Hamilton et al. 1992 

Fragilaria hungarica var. tumida Cleve-Euler Hamilton et al. 1992 

Fragilaria hyemalis Ehrenberg 1856 

Fragilaria incisa (Boyer) Lange-Bertalot Siver et al. 2005 

Fragilaria inflata (Heiden) Hustedt Stoermer & Kreis 1978 

Fragilaria intermedia Grunow Stoermer & Kreis 1978 

Fragilaria intermedia var. continua A. Cleve Stoermer & Kreis 1978 

Fragilaria intermedia var. fallax (Grunow) A. Cleve Stoermer & Kreis 1978 

Fragilaria intermedia var. lanceolata Fusey Stoermer et al. 1999 

Fragilaria interstincta Hohn & Hellerman Hohn & Hellerman 1963 

Fragilaria javanica Hustedt Scherer 1988 

Fragilaria kriegeriana Krasske Stoermer & Kreis 1978 

Fragilaria lanceolata Patrick & Reimer 1966 

Fragilaria lancetmla Schumann Tempere & Peragallo 1908 

Fragilaria lapponica Grunow Stoermer & Kreis 1978 

Fragilaria lapponica var. minuta Cleve Kaczmarska & Rushforth 1983 

Fragilaria lata (Cleve-Euler) Renberg Hamilton et al. 1992 

Fragilaria leptostauron (Ehrenberg) Hustedt Stoermer & Kreis 1978 

Fragilaria leptostauron var. dubia (Grunow) Hustedt Stoermer & Kreis 1978 

Fragilaria leptostauron var. fossilis (Grunow) Rehakova Stoermer & Kreis 1978 

Fragilaria leptostauron var. rhomboides (Grunow) Hustedt Stoermer & Kreis 1978 

Fragilaria levis Ehrenberg Patrick & Reimer 1966 

Fragilaria hnearis Castracane Stoermer & Kreis 1978 

Fragilaria marina var. parva Tempere & Peragallo Tempere & Peragallo 1908 

Fragilaria mazamaensis (Sovereign) Lange-Bertalot Williams & Round 1987 

Fragilaria minuscula Grunow Stoermer & Kreis 1978 

Fragilaria mormonorum (Grunow) Boyer (?) Boyer 1927a 

Fragilaria mutabilis Grunow Stoermer & Kreis 1978 

Fragilaria mutabilis var. intercedens W. Smith Stoermer & Kreis 1978 

Fragilaria neoproducta Lange-Bertalot Siver et al. 2005 

Fragilaria nitida M. Peragallo & Heribaud Tempere & Peragallo 1913 

Fragilaria nitzschioides Grunow Stoermer & Kreis 1978 

Fragilaria oxyptera Ehrenberg Patrick & Reimer 1966 

Fragilaria pantocsekii var. binodis (Pantocsek) A. Cleve Stoermer & Kreis 1978 

Fragilaria paradoxa Ehrenberg Patrick & Reimer 1966 

Fragilaria parasitica (W. Smith) Grunow Stoermer & Kreis 1978 

Fragilaria pectinalis Ehrenberg 1856 

Fragilaria pennsylvanica Morales Morales 2003 

Fragilaria pinnata Ehrenberg Stoermer & Kreis 1978 

Fragilaria pinnata var. acuminata A. Mayer Camburn & Charles 2000 

Fragilaria pinnata var. intercedens (Grunow) Hustedt Stoermer & Kreis 1978 

Fragilaria pinnata var. lancettula (Schumann) Hustedt Stoermer & Kreis 1978 

Fragilaria pinnata var. lancettula f. subcapitata Fusey Camburn & Charles 2000 

Fragilaria pinnata var. subcapitata Frenguelli Reimer 1966 

Fragilaria pinnata var. trigona (Brun & Heribaud) Hustedt Patrick & Reimer 1966 

Fragilaria radians (Kutzing) Williams & Round Stoermer et al. 1999 

Fragilaria rhabdosoma Ehrenberg Patrick & Reimer 1966 

Fragilaria rhodana Hohn & Hellerman Hohn & Hellerman 1963 

Fragilaria robusta Hustedt Patrick & Reimer 1966 

Fragilaria rostrata Ehrenberg Patrick & Reimer 1966 

Fragilaria similis Krasske Grimes & Rushforth 1982 

Fragilaria sinuata M. Peragallo Patrick & Reimer 1966 



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aria smithiana Grunow in Van Heurck Patrick & Reimer 1966 

aria socia (Wallace) Lange-Bertalot Kalinsky 1982 

aria spinosa Skvortzow Stoermer & Kreis 1978 

aria strangulata Patrick & Roberts 1979 

aria striolata Ehrenberg Rushforth & Merkley 1988 

aria sublika Hohn & Hellerman Patrick & Reimer 1966 

aria suboldenburgiana Cambum & Charles Cambum & Charles 2000 

aria synegrotesca Lange-Bertalot Lange-Bertalot 1993 

aria tenera (W. Smith) Lange-Bertalot Kalinsky 1982 

aria turgens Ehrenberg Patrick & Reimer 1966 

aria ulna (Nitzsch) Lange-Bertalot Kalinsky 1982 

aria ulna var. amphirhynchus (Ehrenberg) Kahnsky Kalinsky 1982 

aria ulna var. danica (Kiitzing) Kalinsky Kalinsky 1982 

aria undata W. Smith Boyer 1927a 

aria undata var. lobata Patrick Patrick 1945 

aria undata var. quadrata Hustedt Hohn 1951 

aria vaucheriae (Kiitzing) Peterson Stoermer & Kreis 1978 

aria vaucheriae var. capitellata (Grunow) Patrick Stoermer & Kreis 1978 

aria vaucheriae var. continua A. Cleve Collins & Kalinsky 1977 

aria vaucheriae f. contorta Lowe Dodd 1987 

aria vaucheria var. distans (Grunow) Boye Petersen Patrick & Reimer 1966 

aria vaucheriae var. fallax Grunow Stoermer & Kreis 1978 

aria vaucheriae var. lanceolata A. Mayer Stoermer & Kreis 1978 

aria vaucheriae var. parvula (Kutzing) A. Cleve Stoermer et al. 1999 

aria vaucheriae var. truncata (Greville) Grunow Stoermer & Kreis 1978 

aria venter Ehrenberg 1856 

aria virescens Ralfs Stoermer & Kreis 1978 

aria virescens var. capitata 0strup Stoermer & Kreis 1978 

aria virescens var. clavata Patrick 1968 

aria virescens var. exigua Grunow in Van Heurck Hamilton et al. 1992 

aria virescens var. mesolepta (Ralfs) Schonfeldt Stoermer & Kreis 1978 

aria virescens var. nipha Hohn & Hellerman Hohn & Hellerman 1963 

aria virescens var. oblongella Grunow Stoermer & Kreis 1978 

aria virescens var. producta Lagrstedt Tilden 1894—1909 (#1 19) 

aria virescens var. subsalina Grunow Patrick & Reimer 1966 

ariforma acidobionta (Charles) Williams & Round Williams & Round 1987 

ariforma bicapitata (Mayer) Williams & Round Stoermer et al. 1999 

ariforma constricta (Ehrenberg) Williams & Round Stoermer et al. 1999 

ariforma constricta f. stricta (A. Cleve) Poulin Stoermer et al. 1999 

ariforma constricta f. tetranodis (A. Cleve) Poulin in Hamilton et al Hamilton et al. 1992 

ariforma constricta var. trinodis (Hustedt) Hamilton in Hamilton et al Hamilton et al. 1992 

ariforma hungarica (Pantocsek) Hamilton in Hamilton et al Hamilton et al. 1992 

ariforma hungarica var. tumida (Cleve-Euler) Hamilton in Hamilton et al Hamilton et al. 1992 

ariforma lata (Cleve-Euler) Williams & Round Hamilton et al. 1992 

ariforma virescens (Ralfs) Williams & Round Stoermer et al. 1999 

ariforma virescens var. capitata (0strup) Czamecki Stoermer et al. 1999 

ariforma virescens var. exigua (Grunow) Poulin in Hamilton et al Hamilton et al. 1992 

ariforma virescens var. mesolepta (Rabenhorst) Andresen et al Andresen et al. 2000 

ariforma virescens var. oblongella (Grunow) Andresen et al Andresen et al. 2000 



Fragilariopsis linearis (Castracane) Hustedt Stoermer et al. 1999 

Frustulia asymmetrica (Cleve) Hustedt Patrick & Reimer 1966 

Frustulia bahlsii Edlund & Brant Edlund & Brant 1997 

Frustulia crassiner\'ia (Brebisson) Lange-Bertalot & Krammer Potapova & Charles 2003 

Frustulia disjuncta Lange-Bertalot Lange-Bertalot 2001 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 429 



Name Publication 

Frustulia erifuga Lange-Bertalot & Krammer Stoermer et al. 1999 

Frustulia interposita (Lewis) Cleve Patrick & Reimer 1966 

Frustulia krammeri Lange-Bertalot & Metzeltin in Metzeltin & Lange-Bertalot Siver et al. 2005 

Frustulia longinqua Lange-Bertalot Lange-Bertalot 2001 

Frustulia pelliculosa Brebisson H.L. Smith 1876-1888 (#171) 

Frustulia pseudomagaliesmontana Cambum & Charles Camburn & Charles 2000 

Frustulia rhomboides (Ehrenberg) De Toni Stoermer & Kreis 1978 

Frustuha rhomboides var. amphipleuroides (Grunow) Cleve Stoermer & Kreis 1978 

Frusmlia rhomboides var. capitata (A, Mayer) Patrick Camburn 1982 

Frusmlia rhomboides var. crassinervia (Brebisson) Ross Stoermer & Kreis 1978 

Frustulia rhomboides f. occidentalis Sovereign Patrick & Reimer 1966 

Frusmha rhomboides var. saxonica (Rabenhorst) De Toni Stoenner & Kreis 1978 

Frustulia rhomboides var. saxonica f. capitata A. Mayer Dodd 1987 

Frustulia rhomboides var. saxonica f. undulata Hustedt Dodd 1987 

Frustulia rhomboides f. undulata Hustedt Bateman & Rushforth 1984 

Frustuha rhomboides var. viridula (Brebisson) Cleve Stoermer et al. 1999 

Frustulia saxonica Ehrenberg Stoermer et al. 1999 

Frustulia viridula (Brebisson) De Toni Stoermer & Kreis 1978 

Frustulia vulgaris (Thwaites) De Toni Stoermer & Kreis 1978 

Frustulia vulgaris var. capitata Krasske Stoermer & Kreis 1978 

Frusmlia vveinholdii Hustedt Stoermer & Kreis 1978 

Gallionella aurichalcea Ehrenberg 1856 

Gallionella coarctata Ehrenberg 1856 

Gallionella crenata Ehrenberg 1 856 

Gallionella [sic] crotonensis H.L. Smith 1876-1888 (#678) 

Gallionella distans (Ehrenberg) Stoermer & Kreis 1978 

Gallionella granulata Ehrenberg 1856 

Gallionella laevis Ehrenberg 1856 

Gallionella mar>'landica Ehrenberg 1856 

Gallionella procera Ehrenberg 1856 

Gallionella varians Ehrenberg 1856 

Geissleria acceptata (Hustedt) Lange-Bertalot & Metzeltin Stoermer et al. 1999 

Geissleria aikenensis (Patrick) Torg. & Oliveira Potapova & Charles 2003 

Geissleria declivis (Hustedt) Lange-Bertalot Stoermer et al. 1999 

Geissleria decussis (0strup) Lange-Bertalot & Metzeltin Stoermer et al. 1999 

Geissleria kriegeri (BCrasske) Lange-Bertalot Stoermer et al. 1999 

Geissleria paludosa (Hustedt) Lange-Bertalot & Metzeltin Stoermer et al. 1999 

Geissleria schoenfeldii (Hustedt) Lange-Bertalot & Metzeltin Stoermer et al. 1999 

Geissleria similis (Krasske) Lange-Bertalot & Metzeltin Stoermer et al. 1999 

Geissleria tectissima (Lange-Bertalot) Lange-Bertalot Stoermer et al. 1999 

Geissleria thingvallae (0strupj Metzeltin & Lange-Bertalot Stoermer et al. 1999 

Gloeonema gracile Ehrenberg 1856 

Gloeonema paradoxum Ehrenberg Stoermer & Kreis 1978 

Gloeonema triangulatum Stoermer & Kreis 1978 

Gloeonema virginianum Ehrenberg 1856 

Gomphocymbella ancyli (Cleve) Hustedt Andresen & Stoermer 1978 

Gomphoneis elegans (Grunow) Cleve Boyer 1927b 

Gomphoneis eriense (Grunow) Skvortzow in Skvortzow & Meyer Stoermer & Kreis 1978 

Gomphoneis eriense var. angularis Kociolek & Stoermer Kociolek & Stoermer 1988 

Gomphoneis eriense var. apiculata Stoermer in Reimer Kociolek & Stoermer 1988 

Gomphoneis eriense var. rostrata (M. Schmidt) Skvortzow in Skvortzow & Meyer Kociolek & Stoermer 1988 

Gomphoneis eriense var. variabilis Kociolek & Stoermer Kociolek & Kingston 1999 



430 



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Name Publication 

Gomphoneis geitleri Kociolek & Stoermer Stoermer et al. 1999 

Gomphoneis herculeana (Ehrenberg) Cleve Stoermer & Kreis 1978 

Gomphoneis herculeana var. abundans Kociolek & Stoermer Kociolek & Stoermer 

Gomphoneis herculeana var. clavata Cleve Patrick & Reimer 1975 

Gomphoneis herculeana var. loweii Kociolek & Stoermer Kociolek & Stoermer 

Gomphoneis herculeana var. robusta (Grunow) Cleve Stoermer et al. 1999 

Gomphoneis herculeana var. rostrata Tempere & Peragallo Patrick & Reimer 1975 

Gomphoneis herculeana var. septiceps M. Schmidt Patrick & Reimer 1975 

Gomphoneis linearis Kociolek & Stoermer Kociolek & Stoermer 1986 

Gomphoneis mammilla (Ehrenberg) Cleve Boyer 1927b 

Gomphoneis minuta (Stone) Kociolek & Stoermer Kociolek & Kingston 1999 

Gomphoneis olivaceum (Homemann) Dawson ex Ross & Sims Cambum 1982 

Gomphoneis olivacea var. calcarea (Cleve) Poulin in Pouhn et al Stoermer et al. 1999 

Gomphoneis quadripunctata (0strup) Dawson ex Ross & Sims Stoermer et al. 1999 

Gomphoneis quadripunctata var. cochleariformis Kociolek & Stoermer Stoermer et al. 1999 

Gomphoneis rostrata (Tempere & Peragallo) Kociolek & Stoermer Kociolek & Stoermer 

Gomphoneis rostrata var. valida Kociolek & Stoermer Kociolek & Stoermer 

Gomphoneis scapha M. Schmidt Tempere & Peragallo 

Gomphoneis septa (Moghadam) Kociolek, Stoermer & Bahls Kociolek & Stoermer 

Gomphoneis subherculeana Kociolek & Stoermer Kociolek & Stoermer 

Gomphoneis truUata Kociolek & Stoermer Kociolek & Stoermer 



Gomphonema abbreviatum Agardh Stoermer & Ki^eis 

Gomphonema abbreviatum var. inflata Hustedt Stoermer & Kreis 

Gomphonema acuminatum Ehrenberg Stoermer & Kreis 

Gomphonema acuminatum var. brebissonii (Kutzing) Cleve Stoermer & Kreis 

Gomphonema acuminatum var. capitatum Mayer Collins & Kalinsky 

Gomphonema acuminatum var. clavus (Brebisson) Grunow Cambum 

Gomphonema acuminatum var. coronatum (Ehrenberg) Rabenhorst Stoermer & Kreis 

Gomphonema acuminatum var. elongatum (W. Smith) Carr Cambum 

Gomphonema acuminatum var. intermedia Gmnow Stoermer & Kreis 

Gomphonema acuminatum var. laticeps Ehrenberg Stoermer & Kreis 

Gomphonema acuminatum var. obtusatum (Kiitzing) Gmnow Johansen et al. 

Gomphonema acuminatum var. pusillum Grunow Stoermer & Kreis 

Gomphonema acuminatum var. trigonocephala (Ehrenberg) Gmnow Stoermer & Kreis 

Gomphonema acuminatum var. turns Ehrenberg Stoermer & Kreis 

Gomphonema aequale Gregory Boyer 1 

Gomphonema affme Kutzing Cambum 

Gomphonema affme var. elongatum (Mayer) Millie & Lowe Stoermer et al. 

Gomphonema affme var. insigne (Gregory) Andrews Stoermer & Kreis 

Gomphonema affme f major Gmnow Hohn & Hellerman 

Gomphonema affme var. rhombicum Reichardt Reichardt 

Gomphonema americanum Ehrenberg Stoermer & Kreis 

Gomphonema americobtusatum Reichardt & Lange-Bertalot Reichardt 

Gomphonema angustatum (Kutzing) Rabenhorst Stoermer & Kreis 

Gomphonema angustatum var. angustissima Tempere & Peragallo 

Gomphonema angustatum var. citera (Hohn & Hellerman) Patrick Stoermer et al. 

Gomphonema angustatum var. elongata M. Peragallo in Tempere & Peragallo Tempere & Peragallo 

Gomphonema angustatum var. intermedia Gmnow Cambum 

Gomphonema angustatum var. linearis Hustedt Collins & Kalinsky 

Gomphonema angustatum var. obesa Lauby Gaufm et al. 

Gomphonema angustatum f. major Van Heurck Reimer 

Gomphonema angustatum var. naviculiformis Mayer Collins & Kalinsky 

Gomphonema angustatum var. obesa Lauby Patrick 

Gomphonema angustatum var. obtusatum (Kutzing) Van Heurck Stoermer & Kreis 

Gomphonema angustatum var. productum Gmnow Stoermer & Kreis 

Gomphonema angustatum var. sarcophagus (Gregory) Gmnow Stoermer & Kreis 



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KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 43 1 



Name Publication 

Gomphonema angustatum var. undulata Grunow Stoermer & Kreis 1978 

Gomphonema angustatum f. undulata Grunow Stoermer & Kreis 1978 

Gomphonema apicatum Ehrenberg Patrick & Reimer 1975 

Gomphonema apiculatum Ehrenberg Ehrenberg 1843 

Gomphonema apuncto Wallace Kociolek & Kingston 1999 

Gomphonema argus Ehrenberg Patrick & Reimer 1975 

Gomphonema augur Ehrenberg Stoermer & Kreis 1978 

Gomphonema augur var. gautieri Van Heurck Dodd 1987 

Gomphonema auritum Braun Boyer 1927b 

Gomphonema bohemicum Reichelt & Fricke Stoermer & Kreis 1978 

Gomphonema brasiliense Grunow Stoermer & Kreis 1978 

Gomphonema brasiliense var. rhombiformis Patrick & Roberts 1979 

Gomphonema brebissonii Kutzing Patrick & Reimer 1975 

Gomphonema cambumii Metzeltin & Lange-Bertalot Cambum & Charles 2000 

Gomphonema capitatum Ehrenberg Stoermer & Kreis 1978 

Gomphonema carolinense Hagelstein Hohn & Hellerman 1963 

Gomphonema christensenii Lowe & Kociolek Lowe & Kociolek 1984 

Gomphonema citera Hohn & Hellerman Hohn & Hellerman 1963 

Gomphonema clavatum Ehrenberg Patrick & Reimer 1975 

Gomphonema clavaherculis Ehrenberg 1856 

Gomphonema clevei Fricke Stoermer & Kreis 1978 

Gomphonema commutatum Grunow Patrick 1968 

Gomphonema commutatum var. subramosum Patrick & Reimer 1975 

Gomphonema consector Hohn & Hellerman Stoermer & Kreis 1978 

Gomphonema constrictum Ehrenberg Stoermer & Kreis 1978 

Gomphonema constrictum var. capitata (Ehrenberg) Van Heurck Stoermer & Kreis 1978 

Gomphonema constrictum f. clavata Cleve-Euler Clark & Rushforth 1977 

Gomphonema constrictum var. cuneata A. Schmidt Whitford & Schumacher 1973 

Gomphonema constrictum var. elongata Heribaud & Peragallo Stoermer 1964 

Gomphonema constrictum var. subcapitatum Van Heurck Boyer 1927b 

Gomphonema coronatum Ehrenberg Stoermer & Kreis 1978 

Gomphonema cristatum Ralfs H.L. Smith 1876-1888 (#171) 

Gomphonema cumrhis Hohn & Hellerman Hohn & Hellerman 1963 

Gomphonema cur\atum Kutzing Stoermer & Kreis 1978 

Gomphonema cymbelliclinum Reichardt & Lange-Bertalot Reichardt 1999 

Gomphonema cygnus Ehrenberg Ehrenberg 1843 

Gomphonema dichotomum Kutzing Stoermer & Kreis 1978 

Gomphonema elegans Grunow Rushforth & Merkley 1988 

Gomphonema elongatum W. Smith Tempere & Peragallo 1912 

Gomphonema eriense Grunow Stoermer & Kreis 1978 

Gomphonema exiguum Kutzing Elmore 1922 

Gomphonema exilissima (Grunow) Lange-Bertalot & Reichardt Stoermer et al. 1999 

Gomphonema freesei Lowe & Kociolek Lowe & Kociolek 1984 

Gomphonema geminata (Lyngbye) Agardh Stoermer & Kreis 1978 

Gomphonema germainii Kociolek & Stoermer Stoermer et al. 1999 

Gomphonema gibba Wallace Cambum 1982 

Gomphonema giganteum Ehrenberg Ehrenberg 1852 

Gomphonema glans Ehrenberg Patrick & Reimer 1975 

Gomphonema globiferum Meister Patrick & Reimer 1975 

Gomphonema gracile Ehrenberg Stoermer & Kreis 1978 

Gomphonema gracile var. auritum (A. Braun) Grunow Patrick 1945 

Gomphonema gracile var. cymbelloides Grunow Stoermer & Kreis 1978 

Gomphonema gracile var. insignis Gregory Collins & Kalinsky 1977 

Gomphonema gracile var. intricatiforme Mayer Bateman & Rushforth 1984 

Gomphonema gracile var. janceolata (Kutzing) Cleve Collins & Kalinsky 1977 

Gomphonema gracile var. dichotoma (Kutzing) Grunow Stoermer & Kreis 1978 

Gomphonema gracile f. major (Grunow) O. MUller Hohn 1961 



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Name Publication 

Gomphonema gracile var. naviculacea (W. Smith) Cleve Stoermer & Kreis 1978 

Gomphonema gracile var. naviculoides (W. Smith) Grunow Patrick 1945 

Gomphonema gracile f. parva Van Heurck Reimer 1970 

Gomphonema grovei M. Schmidt Stoermer & Kreis 1978 

Gomphonema grovei var. lingulatum (Hustedt) Lange-Bertalot Stoermer et al. 1999 

Gomphonema grunowii Patrick Cambum 1982 

Gomphonema hedinii Hustedt Patrick & Reimer 1975 

Gomphonema helveticum Brun Stoermer & Kreis 1978 

Gomphonema helveticum var. tenuis Hustedt Stoermer & Kreis 1978 

Gomphonema herculeanum Ehrenberg Stoermer & Kreis 1978 

Gomphonema herculeanum var. robusta Grunow Stoermer & Kreis 1978 

Gomphonema himantaneum Ehrenberg H.L. Smith 1876-1888 (#563) 

Gomphonema hotchkissii Van Landingham Cambum 1982 

Gomphonema innocens Reichardt Reichardt 1999 

Gomphonema insigne Gregory Collins & Kalinsky 1977 

Gomphonema instabilis Hohn & Hellerman Collins & Kalinsky 1977 

Gomphonema intermedium Grunow Tempere & Peragallo 1909 

Gomphonema intricatum Kutzing Stoermer & Kreis 1978 

Gomphonema intricatum var. bohemicum (R. & F.) Cleve-Euler Patrick & Reimer 1975 

Gomphonema intricatum f. pusilla Mayer Collins & Kalinsky 1977 

Gomphonema intricatum var. dichotomum (Kutzing) Grunow Stoermer & Kreis 1978 

Gomphonema intricatum var. fossilis Pantocsek Stoermer & Kreis 1978 

Gomphonema intricatum var. pumila Grunow Stoermer & Kreis 1978 

Gomphonema intricatum var. pulvinatum (Braun) Grunow Cambum 1982 

Gomphonema intricatum var. vibrio (Ehrenberg) Cleve Stoermer & Kreis 1978 

Gomphonema intricatum var. vibrio f. subcapitata A. Mayer Patrick & Reimer 1975 

Gomphonema kobayasii Kociolek & Kingston Kociolek & Kingston 1999 

Gomphonema lanceolatum Agardh Collins & Kalinsky 1977 

Gomphonema lanceolatum var. insignis (Gregory) Cleve Stoermer & Kreis 1978 

Gomphonema laticeps Ehrenberg Stoermer & Kreis 1978 

Gomphonema leptocampum Kociolek & Stoermer Stoermer et al. 1999 

Gomphonema linea Ehrenberg 1 856 

Gomphonema lingulatiforme Lange-Bertalot & Reichardt Potapova & Charles 2003 

Gomphonema lingulatum Hustedt Stoermer & Kreis 

Gomphonema lingulatum var. constricta Hustedt Collins & Kalinsky 

Gomphonema longiceps Ehrenberg Stoermer & Kreis 

Gomphonema longiceps var. montana (Schumann) Cleve Stoermer & Kreis 

Gomphonema longiceps var. subclavata Gmnow Stoermer & Kreis 

Gomphonema longiceps var. subclavata f. gracilis Hustedt Stoermer & Kreis 

Gomphonema longiceps f. gracilis Hustedt Stoermer & Kreis 

Gomphonema longiceps f. suecica Gmnow Hansmann 

Gomphonema longicolle Ehrenberg Patrick & Reimer 

Gomphonema louisiananum Kalinsky Kalinsky 

Gomphonema maclaughlinii Reichardt Reichardt 

Gomphonema mammilla Ehrenberg Ehrenberg 

Gomphonema manubrium Fricke Stoermer & Kreis 

Gomphonema mehleri Cambum Cambum 

Gomphonema me.xicanum Granow Reichardt 

Gomphonema micropus Kutzing Tempere & Peragallo 

Gomphonema minutissima Greville emend. Ehrenberg Patrick & Reimer 

Gomphonema minutum Agardh Kociolek & Kingston 

Gomphonema montanum Schumann Stoermer & Kreis 

Gomphonema montanum var. acuminatum (Peragallo & Heribaud) A. Mayer Patrick & Reimer 

Gomphonema montanum var. media Gmnow Collins & Kalinsky 

Gomphonema montanum var. subclavatum Gmnow Cambum 

Gomphonema montanum var. suecica Gmnow Patrick & Reimer 

Gomphonema mustela Ehrenberg Andresen & Stoermer 



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KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 433 



Name Publication 

Gomphonema mantezumense Czamecki & Blinn Czarnecki 1979 

Gomphonema nasutum Ehrenberg Patrick & Reimer 1975 

Gomphonema novacula Hohn & Hellerman Hohn & Hellerman 1963 

Gomphonema obtusum Ehrenberg 1856 

Gomphonema olivaceoides Hustedt Stoermer & Kreis 1978 

Gomphonema ohvaceoides var. cochleariformis Manguin Stoermer & Kreis 1978 

Gomphonema olivaceoides var. densestriata Foged Stoermer & Kreis 1978 

Gomphonema olivaceoides var. hutchinsoniana Patrick Cambum 1982 

Gomphonema olivaceum (Lyngbye) Kutzing Stoermer & Kreis 1978 

Gomphonema olivaceum var. calcarea (Cleve) Cleve Stoermer & Kreis 1978 

Gomphonema olivaceum var. minutissima Hustedt Stoermer & Kreis 1978 

Gomphonema olivaceum var. olivaceoides (Hustedt) Lange-Bertalot Stoermer et al. 1999 

Gomphonema olivaceum var. tenellum (Kutzing) Cleve Patrick & Reimer 1975 

Gomphonema ohvaceum var. vulgaris (Kutzing) Grunow Patrick & Reimer 1975 

Gomphonema olor Ehrenberg Ehrenberg 1849 

Gomphonema oregonicum Ehrenberg Tempere & Peragallo 1912 

Gomphonema ovamm H.L. smith Patrick & Reimer 1975 

Gomphonema pachycladum Brebisson Collins & Kalinsky 1977 

Gomphonema pala Reichardt Siver et al. 2005 



Gomphonema par%'ulum (Kutzing) Kutzing Stoermer & Kreis 

Gomphonema par\ulum var. aequalis A. Mayer Dodd 

Gomphonema parvoilum var. subelliptica Cleve Stoermer & Kreis 

Gomphonema par\'ulum var. exilissima Grunow Stoermer & Kreis 

Gomphonema parvulum var. lagenula (Kutzing) Frenguelli Stoermer & Kreis 

Gomphonema parvulum var. micropus (Kiitzing) Cleve Stoermer & Kreis 

Gomphonema parvulum var. parvulius Lange-Bertalot & Reichardt Potapova & Charles 

Gomphonema parvulum var. subellipticum Cleve Whitford & Schumacher 

Gomphonema patricki Kociolek & Stoermer Kociolek et al. 

Gomphonema productum Lange-Bertalot & Reichardt Stoermer et al. 

Gomphonema pseudoaugur Lange-Bertalot Stoermer et al. 

Gomphonema pseudopusillum Reichardt Reichardt 

Gomphonema pseudotenellum Lange-Bertalot Stoermer et al. 



Gomphonema puiggarianum Grunow Boyer 1927b 



Gomphonema puiggarianum var. aequatorialis Cleve Cambum 



Gomphonema pumilum (Grunow) Reichardt & Lange-Bertalot Potapova & Charles 2002 

Gomphonema pygmaeum Kociolek & Stoermer Stoermer et al. 1999 

Gomphonema quadripunctatum (0strup) Wislough Stoermer & Kreis 1978 

Gomphonema reimeri (Cambum) Kociolek & Kingston Kociolek & Kingston 1999 

Gomphonema rhombicum Fricke Cambum 1982 

Gomphonema rhombicum f. minor Fricke Sovereign 1958 

Gomphonema robustum Grunow Stoermer & Kreis 1978 

Gomphonema rotundatum Ehrenberg 1856 

Gomphonema sagitta Schumann Prescott & Dillard 1979 

Gomphonema sarcophagus Gregory Stoermer & Kreis 1978 

Gomphonema semiapertum Grunow Stoermer & Kreis 1978 

Gomphonema septata Naegeli Prescott & Dillard 1979 

Gomphonema sepmm Moghadam Patrick & Reimer 1975 

Gomphonema simus Hohn & Hellerman Hohn & Hellerman 1963 

Gomphonema sparsistriatum (O. Muller) Engler Patrick 

Gomphonema sparsistriatum f. maculatum Cambum Cambum 1982 

Gomphonema sphaerophorum Ehrenberg Stoermer & Kreis 1978 

Gomphonema sphaerophorum var. turgidum Ehrenberg Patrick & Reimer 1975 

Gomphonema stoermeri (M. Schmidtj Kociolek & Kingston Kociolek & Kingston 1999 

Gomphonema stonei Reichardt Hohn & Hellerman 1963 

Gomphonema subclavata var. mustela (Ehrenberg) Cleve Stoermer & Kreis 1978 

Gomphonema subclavata f. gracilis (Hustedt) Woodhead & Tweed Stoermer & Kreis 1978 

Gomphonema subclavatum (Grunow) Grunow Stoermer & Kreis 1978 



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Gomphonema subclavatum var. commutatum (Grunow) A. Mayer Patrick & Reimer 1975 

Gomphonema subclavatum var. mexicanum (Grunow) Patrick Cambum 1982 

Gomphonema subclavatum var. mustela (Ehrenberg) Cleve Stoermer et al. 1999 

Gomphonema submehleri Kociolek & Stoermer Stoermer et al. 1999 

Gomphonema subtile Ehrenberg Stoermer & Kreis 1978 

Gomphonema subtilis f. angusta Tempere & Peragallo 1913 

Gomphonema subtile var. sagitta (Schumann) Cleve Stoermer & Kreis 1978 

Gomphonema subventricosum Hustedt Cambum 1982 

Gomphonema superiorensis Kociolek & Stoermer Stoermer et al. 1999 

Gomphonema tackei var. brevistriatum Cambum Cambum et al. 1978 

Gomphonema tenellum Kiitzing Cambum 1982 

Gomphonema tergestinum (Gmnow) Fricke Stoermer & Kreis 1978 

Gomphonema trigonocephalum Ehrenberg Stoermer & JCreis 1978 

Gomphonema tmncatum Ehrenberg Stoermer & Kreis 1978 

Gomphonema truncatum var. capitatum (Ehrenberg) Patrick Cambum 1982 

Gomphonema tmncatum var. cuneatum (Fricke) Cambum Cambum 1982 

Gomphonema truncatum var. elongata (Peragallo & Heribaud) Patrick Patrick & Reimer 1975 

Gomphonema tmncatum var. macilentum Kociolek & Stoermer Stoermer et al. 1999 

Gomphonema truncatum var. turgidum (Ehrenberg) Patrick Stoermer et al. 1999 

Gomphonema tumens Kociolek & Stoermer Kociolek & Stoermer 1991 

Gomphonema turgidum Grunow Stoermer & Kreis 1978 

Gomphonema turris Ehrenberg Stoermer & Kreis 1978 

Gomphonema turritum Ehrenberg 1856 

Gomphonema validum Cleve Patrick & Reimer 1975 

Gomphonema validum var. elongatum Cleve Patrick & Reimer 1975 

Gomphonema variabilis Jurilj Patrick & Reimer 1975 

Gomphonema variostriatum Cambum & Charles Cambum & Charles 2000 

Gomphonema ventricosum Gregory Stoermer & Kreis 

Gomphonema ventricosum var. maxima Cleve Patrick & Reimer 

Gomphonema vibrio Ehrenberg Stoermer & Kreis 

Gomphonema vibrio var. fossile (Pantocsek) R. Ross Stoermer et al. 

Gomphonema vibrio intricatum (Kutzing) R. Ross Stoermer et al. 

Gomphonema vibrio var. pumilum (Grunow in Van Heurck) R. Ross Stoermer et al. 

Gomphonitzschia exigua Sovereign Sovereign 

Gomphosphenia lingulatiformis (Lange-Bertalot & Reichardt) Lange-Bertalot Kociolek & Kingston 

Grammatophora stricta Ehrenberg Rushforth & Merkley 

Gmnowia sinuata Rabenhorst Aubert 

Gyrosigma acuminatum (Kiitzing) Rabenhorst Stoermer & Kreis 

Gyrosigma attenuatum (Kiitzing) Rabenhorst Stoermer & Kreis 

Gyrosigma attenuatum var. hippocampus (W. Smith) Brock Gaufm et al. 

Gyrosigma balticum (Ehrenberg) Rabenhorst Kalinsky 

Gyrosigma delicatuium (W. Smith) Elmore Elmore 

Gyrosigma distortum (W. Smith) Cleve Hohn 

Gyrosigma distortum var. stauroneioides (Grunow) Cleve Patrick & Reimer 

Gyrosigma exilis (Gmnow) Reimer Stoermer & Kreis 

Gyrosigma eximium (Thwaites) Boyer Collins & Kalinsky 

Gyrosigma fasciola (Ehrenberg) Griffen & Henfrey Rushforth & Merkley 

Gyrosigma kuetzingii (Gmnow) Cleve Stoermer & Kreis 

Gyrosigma macmm (W. Smith) Griffen & Henfrey Collins & Kalinsky 

Gyrosigma nodifemm (Gmnow) Reimer Stoermer & Kreis 

Gyrosigma obliquum (Gmnow) Boyer Kalinsky 

Gyrosigma obscumm (W. Smith) Griffen & Henfrey Camburn 



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Name Publication 

Gyrosigma obtusatum (Sullivan & Wormley) Boyer Camburn 1982 

Gyrosigma parkeri (Harrison) Boyer Stoermer & Kreis 1978 

Gyrosigma panulum (Kiitzing) Rabenhorst Collins & Kalinsky 1977 

Gyrosigma peisonis (Grunow) Hustedt Collins & Kalinsky 1977 

G>T0sigma reimeri Sterrenburg Stoermer et al. 1999 

Gyrosigma scalproides (Rabenhorst) Cleve Stoermer & Kreis 1978 

Gyrosigma scalproides var. obliqua (Grunow) Cleve Patrick & Reimer 1966 

Gyrosigma sciotense (Sullivan & Wormley) Cleve Stoermer & Kreis 1978 

Gyrosigma spencerii (Quekett) Griffen & Henfrey Stoermer & Kreis 1978 

Gyrosigma spencerii var. curvula (Grunow) Reimer Stoermer & Kreis 1978 

Gyrosigma spencerii var. nodifera Grunow Stoermer & Kreis 1978 

Gyrosigma strigile (W. Smith) Cleve Stoermer & Kreis 1978 

Gyrosigma temperei Cleve Stoermer & Kreis 1978 

Gyrosigma terr\anum f. fontanum Reimer Patrick & Reimer 1966 

Gyrosigma wormleyi (Sullivant) Boyer Stoermer & Kreis 1978 

Hannaea arcus (^hrenberg) Patrick Stoermer & Kreis 1978 

Hannaea arcus var. amphioxys (Rabenhorst) Patrick Stoermer & Kreis 1978 

Hannaea arcus var. linearis Holmboe Stoermer & Kreis 1978 

Hantzschia amphioxys (Ehrenberg) Grunow Stoermer & Kreis 1978 

Hantzschia amphioxys f. capitata O. Muller Camburn 1982 

Hantzschia amphioxys var. capitata O. Muller Stoermer & Kreis 1978 

Hantzschia amphioxys var. elongata Grunow Tempere & Peragallo 1908 

Hantzschia amphioxys var. intermedia Grunow Clark & Rushforth 1977 

Hantzschia amphioxys var. leptocephala 0strup Clark & Rushforth 1977 

Hantzschia amphioxys var. linearis (O. Muller) Cleve-Euler Rushforth & Merkley 1988 

Hantzschia amphioxys var. major Grunow Hohn 1 95 1 

Hantzschia amphioxys var. vivax (Hantzsch) Grunow Stoermer & Kreis 1978 

Hantzschia distincta-punctate Hustedt Hohn 1961 

Hantzschia elongata Grunow Boyer 1927b 

Hantzschia pseudomarina Hustedt Hohn 1951 

Hantzschia virgata (Roper) Grunow in Grunow & Cleve Stoermer et al. 1999 

Hantzschia virgata var. capitellata Hustedt Stoermer et al. 1999 

Hantzschia vivax var. granulata M. Peragallo in Tempere & Peragallo Tempere & Peragallo 1908 

Himantidium arcus Ehrenberg Kalinsky 1983 

Himantidium bidens Ehrenberg Kalinsky 1983 

Himantidium carinatum Ehrenberg 1856 

Himantidium gracile Ehrenberg 1856 

Himantidium monodon Ehrenberg 1856 

Himantidium parallelum Ehrenberg Kalinsky 1983 

Himantidium pectinale Kutzing Kalinsky 1983 

Himantidium pectinale var. major H.L. Smith 1876-1888 (#193) 

Himantidium temarium Ehrenberg 1856 

Himantidium undulatum W.Smith Kalinsky 1983 

Hippodonta capitata (Ehrenberg) Lange-Bertalot, Metzeltin & Witkowski Stoermer et al. 1999 

Hippodonta costulata (Grunow) Lange-Bertalot, Metzeltin & Witkowski Stoermer et al. 1999 

Hippodonta hungarica (Grunowj Lange-Bertalot, Metzeltin & Witkowski Stoermer et al. 1999 

Hippodonta kaiseri Lange-Bertalot, Metzeltin & Witkowski Lange-Bertalot et al. 1996 

Hippodonta linearis ('0strup) Lange-Bertalot, Metzeltin & Witkowski Stoermer et al. 1999 

Hippodonta lueneburgensis (Grunow) Lange-Bertalot, Metzeltin & Witkowski Stoermer et al. 1999 

Hippodonta subcostulata (Hustedtj Lange-Bertalot, Metzeltin & Witkowski Stoermer et al. 1999 

Homoeocladia acicularis (Kutzing) Kuntze Elmore 1922 

Homoeocladia amphibia (Grunow) Kuntze Elmore 1922 



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Name Publication 

Homoeocladia amphioxys (Ehrenberg) Kuntze Collins & Kalinsky 1977 

Homoeocladia angustata (W. Smith) Kuntze Elmore 1922 

Homoeocladia apiculata (Gregory) Kuntze Elmore 1922 

Homoeocladia arcus (Buhlhein) Kuntze Elmore 1922 

Homoeocladia brebissonii (W. Smith) Kuntze Elmore 1922 

Homoeocladia commutata (Grunow) Kuntze Elmore 1922 

Homoeocladia dissipata (Kiltzing) Kuntze Elmore 1922 

Homoeocladia dubia (W. Smith) Elmore Elmore 1922 

Homoeocladia fasciculata (Grunow) Kuntze Elmore 1922 

Homoeocladia filiformis W. Smith Boyer 1927b 

Homoeocladia frustulum (Kutzing) Kuntze Elmore 1922 

Homoeocladia hungarica (Grunow) Kuntze Elmore 1922 

Homoeocladia intermedia (Hantzsch) Kuntze Elmore 1922 

Homoeocladia lanceolata (W. Smith) Kuntze Elmore 1922 

Homoeocladia linearis (Agardh) Kuntze Elmore 1922 

Homoeocladia obtusa (Lyngbye) Elmore Elmore 1922 

Homoeocladia palea (Kutzing) Kuntze Collins & Kalinsky 1977 

Homoeocladia paxillifer (Miiller) Elmore Elmore 1922 

Homoeocladia punctata (W. Smith) Kuntze Elmore 1922 

Homoeocladia sigma (Kutzing) Kuntze Elmore 1922 

Homoeocladia sigmoidea (Nitzsch) Elmore Stoermer & Kreis 1978 

Homoeocladia spectabilis (Ehrenberg) Kuntze Elmore 1922 

Homoeocladia subtilis (Grunow) Kuntze Elmore 1922 

Homoeocladia tabellaria (Grunow) Kuntze Elmore 1922 

Homoeocladia tryblionella (Hantzsch) Kuntze Elmore 1922 

Homoeocladia umbonata (Ehrenberg) Kuntze Elmore 1922 

Homoeocladia vermicularis (Kutzing) Kuntze Collins & Kalinsky 1977 

Homoeocladia virgata (Roper) Kuntze Elmore 1922 

Homoeocladia vitrea (Norman) Kuntze Elmore 1922 

Hyalodiscus califomicus Collins & Kalinsky 1977 

Hyalodiscus whitneyi Ehrenberg Kaczmarska & Rushforth 1983 

Hyalosynedra laevigata (Grunow) Williams & Round Stoermer et al. 1999 

Hydrosera triquetra Wallich Whitford & Schumacher 1973 

Karayevia clevei (Grunow in Cleve & Grunow) Round & Bukhtiyarova Stoermer et al. 1999 

Karayevia laterostrata (Hustedt) Round & Bukhtiyarova Stoermer et al. 1999 

Kobayasia jaagii (Meister) Lange-Bertalot Stoermer et al. 1999 

Kobayasia subtilissima (Cleve) Lange-Bertalot Stoermer et al. 1999 

Kobayasiella madumensis (J0rgensen) Lange-Bertalot Siver et al. 2005 

Kobayasiella pseudosubtillissima (Manguin) Lange-Bertalot Siver et al. 2005 

Kolbesia kolbei (Hustedt) Round & Bukhtiyarova Stoermer et al. 1999 

Kolbesia ploeonensis (Hustedt) Round & Bukhtiyarova Stoermer et al. 1999 

Krasskella kriegerana (Krasske) Ross & Sims Cambum 1982 

Lemnicola hungarica (Grunow) Round & Basson Johansen et al. 2004 

Licmophora flabellata Whitford 1956 

Licmophora gracilis (Ehrenberg) Grunow Rushforth & Merkley 1988 

Licmophora tincta (Agardh) Grunow Whitford & Schumacher 1973 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 437 



Name Publication 

Luticola cohnii (Hilse) D.G. Mann in Round et al Stoermer et al. 1999 

Luticola goeppertiana (Bleisch) D.G Mann in Round et al Johansen et al. 2004 

Luticola mutica (Kiitzing) D.G Mann in Round et al Stoermer et al. 1999 

Luticola muticoides (Hustedt) D.G Mann in Round et al Stoermer et al. 1999 

Luticola muticopsis (Van Heurck) D.G. Mann in Round et al Stoermer et al. 1999 

Luticola naviculoides Johansen in Johansen et al Johansen et al. 2004 

Luticola nivalis (Ehrenberg) Andresen et al Andresen et al. 2000 

Luticola saxophila (Bock ex Hustedt) D.G Mann Gaiser & Johansen 2000 

Luticola stigma (Patrick) Johansen in Johansen et al Johansen et al. 2004 

Luticola terminata (Hustedt) Johansen in Johansen et al Johansen et al. 2004 

Luticola terminata var. rostrata (Krasske) Johansen in Johansen et al Johansen et al. 2004 

Luticola undulata (Hilse) Andresen et al Andresen et al. 2000 



Lysigonium crenulatum (Kiitzing) Kuntze Elmore 

Lysigonium distans (Kiitzing) Kuntze Elmore 

Lysigonium granulata (Ehrenberg) Kuntze Stoermer & Kxeis 

Lysigonium varians (Agardh) De Toni Stoermer & Kreis 

Martyana martyi (Heribaud) Round in Round et al Stoermer et al. 

Mastogloia angusta Hustedt Hustedt 

Mastogloia aquilegiae Grunow Kaczmarska & Rushforth 

Mastogloia braunii Grunow Stoermer & Kreis 

Mastogloia crucicula (Grunow) Cleve Kalinsky 

Mastogloia dansei Thwaites Hohn 

Mastogloia doddii Stoermer ex Hungerford Reimer 

Mastogloia dubia Kiitzing Kalinsky 

Mastogloia elhptica (Agardh) Schonfeldt Elmore 

Mastogloia elliptica var. danseii (Thwaites) Cleve Patrick & Reimer 

Mastogloia exigua Kutzing Tempere & Peragallo 

Mastogloia grevillei W. Smith Stoermer & Kreis 

Mastogloia jurgensii Ag Kalinsky 

Mastogloia lacustris (Grunow) Van Heurck Stoermer & Kreis 



Mastogloia lanceolata Kutzing Tilden 1894-1909 (#248) 

Mastogloia pumila (Grunow) Cleve Kalinsky 1983 

Mastogloia smithii Thwaites Stoermer & Kreis 1978 

Mastogloia smithii var. amphicephala Grunow Stoermer & Kreis 1978 

Mastogloia smithii var. lacustris Grunow Stoermer & Kreis 1978 

Melosira agassizzi Ostenfeld Stoermer & Kreis 1978 

Melosira agassizzi var. malayensis Hustedt Stoermer & Kreis 1978 

Melosira ambigua (Grunow) O. Muller Stoermer & Kreis 1978 

Melosira arenaria Moore Stoermer & Kreis 1978 

Melosira arenti (Kolbe) Nagumo & Kobayasi Hamilton et al. 1992 

Melosira binderana Kutzing Stoermer & Kreis 1978 

Melosira borreri Greville Whitford 1956 

Melosira canadensis Hustedt Stoermer & Kreis 1978 

Melosira crenulata (Ehrenberg) Kiitzing Stoermer & Kreis 1978 

Melosira crenulata var. tenuis Kutzing Stoermer & Kreis 1978 

Melosira crotonensis (J.W. Bailey) H.L. Smith Stoermer & Kreis 1978 

Melosira dendroteres (Rabenhorst) R. Ross Stoermer et al. 1999 

Melosira dickiei (Thwaites) Kiitzing Stoermer et al. 1999 

Melosira distans (Ehrenberg) Kutzing Stoermer & Kreis 1978 

Melosira distans var. africana Miiller Hamilton et al. 1992 

Melosira distans var. alpigena Grunow Stoermer & Kreis 1978 

Melosira distans var. humilis Cleve-Euler Camburn & Kingston 1986 

Melosira distans var. limnetica O. Miiller Stoermer et al. 1999 



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Melosira distans var. nivalis (W. Smith) Kirchner Cambum & Kingston 

Melosira distans var. nivaloides Camburn Cambum & Kingston 

Melosira distans var. pfaffiana (Reinsch) Grunow Rushforth & Merkley 

Melosira distans var. tenella (Nygaard) Florin Cambum & Kingston 

Melosira dubia Kiitzing Rushforth et al. 

Melosira epidendron (Ehrenberg) Boyer Boyer 1927a 

Melosira granulata (Ehrenberg) Ralfs Stoermer & Kreis 

Melosira granulata var. angustissima O. Miiller Stoermer & Kreis 

Melosira granulata var. angustissima f. curvata Gmnow Stoermer & Kreis 

Melosira granulata var. angustissima f. spiralis Muller Whitford & Schumacher 

Melosira granulata f. curvata Gmnow Stoermer et al. 

Melosira granulata var. muzzanensis (Meister) Bethge Stoermer & Kreis 

Melosira granulata var. procera (Ehrenberg) Gmnow Cambum 

Melosira granulata f. spiralis Gmnow Stoermer & Kreis 

Melosira herzogii Lemmermann Cambum & Kingston 

Melosira islandica O. Muller Stoermer & Kreis 

Melosira islandica subsp. helvetica O. Muller Stoermer & Kreis 

Melosira italica (Ehrenberg) Kiitzing Stoermer & Kreis 

Melosira italica var. multistriata Patrick Hohn 

Melosria italica var. granulata Gmnow Whitford & Schumacher 

Melosria italica var. subarctica O. Muller Gaufm et al. 

Melosira italica var. tenuis (Gmnow). O. Muller Stoermer et al. 

Melosira italica var. tenuissima (Gmnow) O. Muller Stoermer & Kreis 

Melosira italica var. valida Gmnow Stoermer & Kreis 

Melosira italica subsp. subarctica O. Muller Stoermer & Kreis 

Melosira italica subsp. subarctica f. tenuissima (Gmnow) Cambum Cambum & Kingston 

Melosira juergensii C.A. Agardh Dodd 

Melosira lacustris H.H. Chase Stoermer & Kreis 

Melosira laevis (Ehrenberg) Gmnow Hohn 

Melosira lirata (Ehrenberg) Kiitzing Cambum & Kingston 

Melosira lirata f. biseriata (Gmnow) Cambum Cambum & Kingston 

Melosira lirata var. lacustris Gmnow Cambum & Kingston 

Melosira longispina Hustedt Stoermer & Kreis 

Melosira moniliformis (O.F. Muller) Agardh J.W. Bailey 

Melosira nummuloides (Dillwyn) Agardh • Kalinsky 

Melosira nygaardii Camburn Cambum & Kingston 

Melosira pensacolae A. Schmidt Sovereign 

Melosira perglabra 0stmp Cambum & Kingston 

Melosira perglabra var. floriniae Cambum Camburn & Kingston 

Melosira pseudoamericana Cambum Cambum & Kingston 

Melosira punctata W. Smith Cleve & Moller 

Melosira roseana Rabenhorst Stoermer & Kreis 

Melosira roeseana var. epidendron (Ehrenberg) Gmnow Patrick 

Melosira scalaris Gmnow Tempere & Peragallo 

Melosira semilaevis Gmnow Tempere & Peragallo 

Melosira solida Eulenstein Tempere & Peragallo 

Melosira tenuis Kiitzing Tempere & Peragallo 

Melosira tenuis var. ambigua Gmnow Cleve & Moller 

Melosira tenuissima Gmnow Tempere & Peragallo 

Melosira undulata (Ehrenberg) Kiitzing Stoermer & Kreis 

Melosira undulata var. debilis Tempere & Peragallo 

Melosira undulata var. normanii Amott Stoermer & Kreis 

Melosira varians Agardh Stoermer & Kreis 

Melosira varennamm M. Peragallo Tempere & Peragallo 

Meridion alansmithii Brant Brant 2003 

Meridion anceps (Ehrenberg) Williams Stoermer et al. 1999 



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Name 



Publication 



Meridion circulare (Greville) Agardh Stoermer & Kreis 1978 

Meridion circulare var. constrictum (Ralfs) Van Heurck Stoermer & Kreis 1978 

Meridion constricutm Ralfs Stoermer & Kreis 1978 

Meridion constrictimi var. elongata Tempere & Peragallo Tempere & Peragallo 1908 

Meridion constrictum var. zinkenii Grunow Tempere & Peragallo 1908 

Meridion hohii Rhode Rhode et al. 2001 

Meridion lineare (H.L. Smith) D.M. Williams Williams 1985 

Meridion intermedium H.L. Smith Stoermer & Kreis 1978 

Meridion intermedium var. constrictum H.L. Smith Stoermer & Kreis 1978 

Meridion vemale Agardh Stoermer & Kreis 1978 

Meridion zenkenii Kutzing Tempere & Peragallo 1912 

Microcostatus krasskei (Hustedt) Johansen & Sray Johansen & Sray 1997 

Microsiphona potamos Weber Stoermer & Kreis 1978 

Muelleria gibbula (Cleve) Spaulding & Stoermer Spaulding et al. 1999 

Muelleria terrestris (Petersen) Spaulding & Stoermer Spaulding et al. 1999 



Navicula abaujensis (Paut.) Beitr Myers 1989b 

Navicula abiskoensis Hustedt Collins & Kalinsky 1977 

Navicula aboensis (Cleve) Hustedt Stoermer & Kreis 1978 

Navicula absoluta Hustedt Stoermer & Kreis 1978 

Navicula abunda Hustedt Kalinsky 1983 

Navicula acceptata Hustedt Stoermer & Kreis 1978 

Navicula accomoda Hustedt Stoermer & Kreis 1978 

Navicula achthera Hohn & Hellerman Hohn & Hellerman 1963 

Navicula acrosphaeria (Brebisson) Kutzing Elmore 1922 

Navicula acrosphaeria var. brevis Tempere & Peragallo 1909 

Navicula acrosphaeria var. dilata Tempere & Peragallo Tempere & Peragallo 1908 

Navicula acrosphaeria var. laevis A. Schmidt Tempere & Peragallo 1908 

Navicula acrosphaeria var. minor M. Peragallo & Heribaud Tempere & Peragallo 1908 

Navicula admenda Hohn & Helleerman Patrick & Reimer 1966 

Navicula adnata Hustedt Patrick & Reimer 1966 

Navicula adumbrata Hohn & Hellerman Patrick & Reimer 1966 

Navicula adversa Krasske Hohn & Hellerman 1963 

Navicula aequalis (Ehrenberg) Kutzing Patrick & Reimer 1966 

Navicula aestuarii Cleve Tempere & Peragallo 1911 

Navicula affmis Ehrenberg Stoermer & Kreis 1978 

Navicula affme f. maxima Ehrenberg Tempere & Peragallo 1908 

Navicula agma Hohn & Hellerman Hohn & Hellerman 1963 

Navicula agnita Hustedt Stoermer et al. 1999 

Navicula agrestis Hustedt Hohn & Hellerman 1963 

Navicula aikenenses Patrick Stoermer & Kreis 1978 

Navicula alea Hohn & Hellerman Hohn & Hellerman 1963 

Navicula algor Hohn & Hellerman Patrick & Reimer 1966 

Navicula alpina W. Smith Aubert 1895 

Navicula alpina var. elongata M. Peragallo in Tempere & Peragallo Tempere & Peragallo 1908 

Navicula ambigua Ehrenberg Stoermer & Kreis 1978 

Navicula ambigua var. craticularis Ehrenberg Stoermer & Kreis 1978 

Navicula americana Ehrenberg Stoermer & Kreis 1978 

Navicula americana var. alastos Hohn & Hellerman Stoermer et al. 1999 

Navicula americana var. bacillaris Peragallo & Heribaud Stoermer 1964 

Navicula americana var. minor Peragallo & Heribaud Boyer 1927b 

Navicula americana var. moesta Tempere & Peragallo Patrick & Reimer 1966 

Navicula ammophila var. flanatica (Grunow) Cleve Patrick & Reimer 1966 

Navicula amnicola Hohn & Hellerman Hohn & Hellerman 1963 



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Name Publication 

Navicula amphibola Cleve Boyer 1927b 

Navicula amphibola var. perrieri Peragallo & Heribaud Stoermer & Kreis 1978 

Navicula amphibola var. polymorpha Fusey Stoermer & Kreis 1978 

Navicula amphibola var. stauroneiformis Tempere & Peragallo 1909 

Navicula amphiceros Kiitzing Stoermer & Kreis 1978 

Navicula amphigomphus Ehrenberg Stoermer & Kreis 1978 

Navicula amphilepta Ehrenberg 1856 

Navicula amphioxys Ehrenberg Patrick & Reimer 1966 

Navicula amphirhynchus Ehrenberg Kalinsky 1983 

Navicula amphisbaena Bory Elmore 1922 

Navicula amphisphenia Ehrenberg Kalinsky 1983 

Navicula ampliata Ehrenberg Tempere & Peragallo 1909 

Navicula amydalina Hustedt Stoermer et al. 1999 

Navicula anatis Hohn & Hellerman Hohn & Hellerman 1963 

Navicula anglica Ralfs Stoermer & Kreis 1978 

Navicula anglica var lapponica Cleve-Euler Clark & Rushforth 1977 

Navicula anglica var. signata Hustedt Stoermer & Kreis 1978 

Navicula anglica var. subsalsa (Grunow) Cleve Stoermer & Kreis 1978 

Navicula angusta Grunow Cambum 1982 

Navicula anglica var. subsalsa (Grunow) Cleve Stoermer & Kreis 1978 

Navicula angustata W. Smith Stoermer & Kreis 1978 

Navicula annexa Hustedt Dodd 1987 

Navicula antinitescens M. Peragallo in Tempere & Peragallo Tempere & Peragallo 1908 

Navicula apiculata Brebisson Patrick & Reimer 1966 

Navicula appendiculata Kiitzing Stoermer & Kreis 1978 

Navicula arata Grunow Boyer 1927b 

Navicula arctissima A. Schmidt Tempere & Peragallo 1913 

Navicula arenaria Donkin Patrick & Reimer 1966 

Navicula arenula Hohn & Hellerman Patrick & Reimer 1966 

Navicula argutiola Hohn & Hellerman Hohn & Hellerman 1963 

Navicula arvensis Hustedt Stoermer & Kreis 1978 

Navicula arvema M. Peragallo & Heribaud Patrick & Reimer 1966 

Navicula aspera Ehrenberg Tilden 1894-1909 (#367) 

Navicula asymbasia Hohn & Hellerman Hohn & Hellerman 1963 

Navicula aszellus Weinhold ex Hustedt Reimer 1970 

Navicual atomoides Grunow in V.H Elmore 1922 

Navicula atomus (Kiitzing) Grunow Stoermer & Kreis 1978 

Navicula atomus var. permitis (Hustedt) Lange-Bertalot Stoermer et al. 1999 

Navicula atomus var. recondita (Hustedt) Lange-Bertalot Stoermer et al. 1999 

Navicula auriculata Hustedt Cambum 1982 

Navicula aurora Sovereign Stoermer & Kreis 1978 

Navicula avenacea Brebisson in Grunow Hohn 1961 

Navicula bacillaris Gregory Stoermer & Kreis 1978 

Navicula bacillariformis Grunow Stoermer & Kreis 1978 

Navicula bacilloides Hustedt Fee 1967 

Navicula bacillum Ehrenberg Stoermer & Kreis 1978 

Navicula bacillum var. lepida (Gregory) Cleve Rushforth & Merkley 1988 

Navicula bacula Hohn & Hellerman Hohn & Hellerman 1963 

Navicula balcanica Hustedt Stoermer & Kreis 1978 

Navicula bastianii M. Pergallo in Tempere & Peragallo Boyer 1927b 

Navicula bdesma Hohn Patrick & Reimer 1966 

Navicula begeri Krasske Stoermer & Kreis 1978 

Navicual belliatula Archibald Dodd 1987 

Navicula bergenensis Hohn Patrick & Reimer 1966 

Navicula bicapitata Lagerstedt Stoermer & Kreis 1978 

Navicula bicapitata var. hybrida Grunow Tempere & Peragallo 1908 

Navicula bicapitellata Hustedt Stoermer & Kreis 1978 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 441 



Name Publication 



Navicula bicephala Hustedt Stoermer & Kreis 

Navicula biceps Ehrenberg Collins & Kalinsky 

Navicula biconica Patrick Patrick 

Navicula bicontracta 0strup Stoermer & Kreis 

Na%icula bievexa Hohn Patrick & Reimer 

Navicula binodis Ehrenberg Stoermer & Kreis 

Naxicula birhis Hohn Patrick & Reimer 

Navicula bisulcata Lagerstedt Elmore 

Navicula bita Hohn Patrick & Reimer 

Navicula bogotensis var. ininterrupta M. Peragallo in Tempere & Peragallo Tempere & Peragallo 

Navicula bogotensis var. undulata M. Peragallo in Tempere & Peragallo Tempere & Peragallo 

Na^•icula bohemica Ehrenberg Rushforth & Merkley 

Navicula borealis (Ehrenberg) Kiitzing Elmore 

Navicula bottnica Grunow Bateman & Rushforth 

Navicula braunii Grunow Elmore 

Navicula braunii var. interrupta Tempere & Peragallo 

Navicula brebissonii Kiitzing Stoermer & Kreis 

Navicula brebissonii var. curta Tempere & Peragallo 

Navicula brekkaensis J.B. Petersen Bateman & Rushforth 

Navicula bremensis Hustedt Dixit & Smol 

Navicula brevis Gregory Collins & Kalinsky 

Navicula brevissima Hustedt Gaiser & Johansen 

Navicula brockmannii Hustedt Dodd 

Navicula bryophila 0strup Patrick & Reimer 

Navicula bryophila Peterson Stoermer & Kreis 

Navicula buccella Hohn & Hellerman Patrick & Reimer 

Navicula caduca Hustedt Hohn & Hellerman 

Navicula campylogramma Ehrenberg Stoermer & Kreis 

Navicula canalis Patrick Stoermer & Kreis 

Navicula cancellata var. retusa (Brebisson) Cleve Patrick & Reimer 

Navicula canoris Hohn & Hellerman Hohn & Hellerman 

Navicula capitata Ehrenberg Stoermer & Kreis 

Navicula capitata var. hungarica (Grunow) Ross Stoermer & Kreis 

Navicula capitata var. hnearis (0strup) Stoermer & Kreis Stoermer & Kreis 

Navicula capitata var. luneburgensis (Grunow) Patrick Stoermer & Kreis 

Navicula capitata var. lueneburgensis f. elegans 0strup Stoermer et al. 

Navicula capitatoradiata Germain Yearsley et al. 

Navicula capsa Hohn Stoermer & Kreis 

Navicula caractacus Hohn & Hellerman Hohn & Hellerman 

Navicula cardinalis Ehrenberg Stoermer & Kreis 

Navicula cari Ehrenberg Stoermer & Kreis 

Navicula cari var. angusta Grunow Stoermer & Kreis 

Navicula carminata Patrick 

Navicula camiolensis Hustedt Hohn & Hellerman 

Navicula caroliniana Patrick Stoermer & Kreis 

Navicula carolinensis Ehrenberg Patrick & Reimer 

Navicula cascadensis Sovereign Stoermer & Kreis 

Navicula caterva Hohn & Hellerman Hohn & Hellerman 

Navicula cemeutia Hohn Patrick & Reimer 

Navicula cerla Hustedt Stoermer & Kreis 

Navicula charlatii M. Peragallo Dodd 

Navicula charlatii f. simplex Hustedt Loescher 

Navicula cincta (Ehrenberg) Ralfs Stoermer & Kreis 

Navicula cincta var. angusta (Grunow) Cleve Patrick 

Navicula cincta var. heufleri Tempere & Peragallo 

Navicula cincta var. leptocephala Brebisson ex Van Heurck Collins & Kalinsky 

Navicula cincta var. minuta Grunow Patrick & Reimer 



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442 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Name Publication 

Navicula cincta var. rostrata Reimer Collins & Kalinsky 

Navicula cinna Hohn & Hellerman Hohn & Hellerman 

Navicula circumtexta Meister Stoermer & Kreis 

Navicula citrus Krasske Stoermer & Kreis 

Navicula damans Hustedt Stoermer & Kreis 

Navicula dementis Grunow Stoermer & Kreis 

Navicula dementis var. linearis Brander Stoermer & Kreis 

Navicula dementis var quadristigmata Manguin Stoermer & Kreis 

Navicula clementoides Hustedt Stoermer et al. 

Navicula cocconeiformis Gregory Stoermer & Kreis 

Navicula cocconeis (Ehrenberg) De Toni Patrick & Reimer 

Navicula Columbiana Hustedt Hustedt 

Navicula columnaris Ehrenberg Tempere & Peragallo 

Navicula commutata Grunow Tempere & Peragallo 

Navicula complanatula Hustedt Hustedt 

Navicula concava Patrick Patrick 

Navicula confervacea (Kutzing) Grunow Stoermer & Kreis 

Navicula confervacea var. peregrina (W. Smith) Grunow Stoermer & Kreis 

Navicula constans Hustedt Clark & Rushforth 

Navicula constans var. symmetrica Hustedt Stoermer & Kreis 

Navicula contempta Hustedt Collins & Kalinsky 

Navicula contenta Grunow Stoermer & Kreis 

Navicula contenta f. biceps Patrick 

Navicula contenta f. parallela (Petersen) Hustedt Rushforth & Merkley 

Navicula contenta var. biceps (Amott) Grunow Stoermer & Kreis 1978 

Navicula contortula Sovereign Patrick & Reimer 1966 

Navicula contraria Patrick Stoermer & Kreis 1978 

Navicula convergens Patrick Patrick 1959 

Navicula costata Ehrenberg Tempere & Peragallo 1909 

Navicula costulata Cleve & Grunow Stoermer & Kreis 1978 

Navicula costuloides Skvortzow Stoermer et al. 1999 

Navicula cremorne Hohn & Hellerman Hohn & Hellerman 1963 

Navicula cristula Hohn Patrick & Reimer 1966 

Navicula crucialis (O. Muller) Frenguelli Kociolek & Herbst 1992 

Navicula crucicula (W. Smith) Donkin Stoermer & Kreis 1978 

Navicula creuzburgensis var multistriata Patrick Patrick 1959 

Navicula cryptocefalsa Lange-Bertalot Stoermer et al. 1999 

Navicula cryptocephala Kutzing Stoermer & Kreis 1978 

Navicula cryptocephala var exilis (Kutzing) Grunow Cambum 1982 

Navicula cryptocephala var. intermedia Van Heurck Stoermer & Kreis 1978 

Navicula cryptocephala var. lancettula (Schumann) Grunow Stoermer & Kreis 1978 

Navicula cryptocephala f. minuta Boye Peterson Hohn & Hellerman 1963 

Navicula cryptocephala var. pumila Grunow Patrick 1945 

Navicula cryptocephala f. terrestris Lund Collins & Kalinsky 1977 

Navicula cryptocephala var. veneta (Kiitzing) Rabenhorst Stoermer & Kreis 1978 

Navicula cryptocephaloides Hustedt Stoermer & Kreis 1978 

Navicula cryptogaster Lowe Lowe 1972-1973 

Navicula cryptonella Lange-Bertalot Stoermer et al. 1999 

Navicula cryptotonella Lange-Bertalot Potapova & Charles 2002 

Navicula cuspidata (Kutzing) Kutzing Stoermer & Kreis 1978 

Navicula cuspidata var. ambigua (Ehrenberg) Cleve Boyer 1927b 

Navicula cuspidata var. danaica Grunow in Cleve Patrick & Reimer 

Navicula cuspidata var. heribaudi Peragallo Patrick & Reimer 

Navicula cuspidata var. major Meister Boyer 1927b 

Navicula cuspidata var. obtusa Patrick Patrick & Reimer 

Navicula cynthia A. Schmidt Collins & Kalinsky 

Navicula cyrpinus W. Smith Tempere & Peragallo 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 443 



Navicula dailyi Reimer Patrick & Reimer 

Navicula dariana A. Schmidt Tempere & Peragallo 

Navicula declivis Hustedt Stoermer et al. 

Navicula decurrens (Ehrenberg) Kutzing Elmore 

Navicula decussis 0strup Stoermer & Kreis 

Navicula demerara Gmnow ex Cleve Patrick & Reimer 

Navicula demissa Hustedt Drum 

Navicula denestriata Hustedt Stoermer & Kreis 

Navicula detenta Hustedt Stoermer et al. 

Navicula diagonalis Patrick & Reimer 

Navicula dicephala Ehrenberg Stoermer & Kreis 

Navicula dicephala var. abiskonensis (Hustedt) A. Cleve Stoermer & Kreis 

Navicula dicephala var. elginensis (Gregory) Cleve Stoermer & Kreis 

Navicula dicephala var. lata M. Peragallo in Tempere & Peragallo Tempere & Peragallo 

Navicula dicephala var. subcapitata Grunow Patrick & Reimer 

Navicula dibola Hohn Patrick & Reimer 



Name Publication 

Navicula dactylus Ehrenberg Aubert Le Diatomiste #20 

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Navicula difficillima Hustedt Cambum & Charles 2000 

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Navicula difficillimoides Hustedt Grimes & Rushforth 

Navicula digito-radiata (Gregory) Ralfs Collins & Kalinsky 

Navicula digimlus Hustedt Bateman & Rushforth 

Navicula dilatata Ehrenberg Ehrenberg 

Navicula diluviana Krasske Stoermer & Kreis 

Navicula diserta Hustedt Kalinsky 

Navicula disjuncta Hustedt Hamilton et al. 

Navicula dismutica Hustedt Loescher 

Navicula disputans Patrick Stoermer & Kreis 

Navicula dissipata Hustedt Stoermer & Kreis 

Navicula distinctastriata Hohn & Hellerman Stoermer & Kreis 

Navicula divergens (W. Smith) Ralfs Stoermer & Kreis 

Navicula divergens var. bacillaris M.Peragallo in Tempere & Peragallo Tempere & Peragallo 

Navicula dubia W. Smith Stoermer & Kreis 

Navicula dubia var. acuminata Tempere & Peragallo Tempere & Peragallo 

Navicula dulcis Patrick Stoermer & Kreis 

Navicula duomedia Patrick Patrick 

Navicula dystrophica Patrick Patrick 

Navicula ebor Hohn & Hellerman Hohn & Hellerman 

Navicula eiowana Ehrenberg Patrick & Reimer 

Navicula elaphros Hohn & Hellerman Hohn & Hellerman 

Navicula elata Gandhi Collins & Kalinsky 

Navicula elegans W. Smith Stoermer & Kreis 

Navicula elegans var. cuspidata Cleve Hohn 

Navicula elegantissima M. Peragallo in Tempere & Peragallo Tempere & Peragallo 

Navicula elginensis (Gregory) Ralfs Stoermer & Kreis 

Navicula elginensis f . abiskoensis Hustedt Clark & Rushforth 

Navicula elginensis var. lata (M. Peragallo) Patrick Stoermer & Kreis 

Navicula elginensis var. neglecta (Krasske) Patrick Cambum 

Navicula elginensis var. rostrata (A. Mayer) Patrick Stoermer & Kreis 

Navicula elginensis var. subcapitata Grunow Clark & Rushforth 

Navicula elliptica Kutzing Stoermer & Kreis 

Navicula elliptica var. minutissima Grunow Stoermer & Kreis 

Navicula elliptica var. ostracodarum Tempere & Peragallo 

Navicula elmorei Patrick Patrick & Reimer 

Navicula elongata Ehrenberg 

Navicula entomon Ehrenberg Collins & Kalinsky 

Navicula eponka Hohn Patrick & Reimer 

Navicula erifuga Lange-Bertalot Johansen et al. 2004 



444 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Name Publication 



Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu^ 
Navicu^ 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu^ 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 



a esox Ehrenberg Ehrenberg 1856 

a evexa Sovereign Patrick & Reimer 1966 

a excelsa Krasske Stoermer & Kreis 1978 

a exigua (Gregory) Grunow Stoermer & Kreis 1978 

a exigua var. capitata Patrick Stoermer & Kreis 1978 

a exiguaformis Hustedt Stoermer & Kreis 1978 

a exibuoides Hustedt Stoermer & Kreis 1978 

a exilis KUtzing Stoermer & Kreis 1978 

a exilissima Grunow Collins & Kalinsky 1977 

a explanata Hustedt Stoermer & Kreis 1978 

a exselsa Krasske Rushforth & Merkley 1988 

a exsul A. Scliniidt Patrick & Reimer 1966 

a falaisensis Grunow Boyer 1927b 

a farta Hustedt Stoermer & Kreis 1978 

a fasciata Lagerstedt Tempere & Peragallo 1908 

a festiva Krasske Stoermer & Kreis 1978 

a feuerbomi Hustedt Kaczmarska & Rushforth 1983 

a fmnica Cleve Tempere & Peragallo 1909 

a firma Kutzing & Grunow Aubert 1 895 

a firma var. amphigomphus Ehrenberg Cleve & MoUer 1 879 

a firma var. dilatata Cleve & Moller 1878 

a firma var. iridis Cleve & Moller 1 879 

a firma var. subampliata Tempere & Peragallo 1912 

a firma var. tumescens Grunow Cleve & Moller 1878 

a flanatica Grunow Kalinsky 1983 

a flavasinus Moghadam Prescott & Dillard 1979 

a flexuosa Ehrenberg 1856 

a flexuosa var. cuneata Tempere & Peragallo Tempere & Peragallo 1909 

a fluens Hustedt Dodd 1987 

a tluminitica Cambum Cambum 1982 

a forcipata Greville Patrick & Reimer 1966 

a formica Ehrenberg Cleve & Moller 1878 

a fossalis Krasske Collins & Kalinsky 1977 

a fracta Hustedt Stoermer & Kreis 1978 

a fragilarioides Krasske Bateman & Rushforth 

a friesneri Reimer Patrick & Reimer 1966 

a fritschii Lund Patrick & Reimer 1966 

a frugalis Hustedt Hohn & Hellerman 1963 

a fulva (Nitzsch) Ehrenberg Elmore 1922 

a fusidium Ehrenberg Kalinsky 1983 

a gallica (W. Smith) Van Heurck Johansen et al 1983 

a gallica var. nitzschioides Grunow Patrick & Reimer 1966 

a gallica var. perpusilla (Grunow) Lange-Bertalot Cambum & Charles 2000 

a gastriformis Hustedt Stoermer & Kreis 1978 

a gastrum (Ehrenberg) Kutzing Stoermer & Kreis 1978 

a gastrum f. maxima Tempere & Peragallo Tempere & Peragallo 1909 

a gastrum var. exigua (Gregory) Grunow Patrick & Reimer 1966 

a gastrum var. signata Hustedt Stoermer & Kreis 1978 

a gaufinii Moghadam Prescott & Dillard 1979 

a genovefea Fusey Hohn & Hellerman 1963 

a gentilis Donkin Tempere & Peragallo 1908 

a germanii Wallich Hohn 1961 

a germainii Wallace Patrick 1968 

a gibba (Ehrenberg) Kutzing Stoermer & Kreis 1978 

a gibba van hyalina Tempere & Peragallo 1909 

a gibhosa Hustedt Stoermer & Kreis 1 978 

a gibbula Cleve Patrick & Reimer 1966 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 445 



Name 

Navicula 

Navicula 

Navicula 

Navicula 

Navicula 

Navicula 

Navicula 

Navicula 

Navicu 

Navicu] 

Navicu^ 

Navicu 

Navicu 

Navicu^ 

Navicu^ 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu 

Navicu^ 

Navicu 

Navicu' 

Navicu 

Navicu 

Navicu 



Publication 

gigas Ehrenberg Tempere & Peragallo 1908 

globiceps Gregory Stoermer & Kreis 1978 

globosa Meister Stoermer & Kreis 1978 

globulifera Hustedt Patrick 1945 

goeppertiana (Bleisch) H.L. Smith Stoermer et al. 1999 

goersii Bahls Bahls 1983 

gottlandica Grunow Stoermer & Kreis 1978 

gracilis Ehrenberg Stoermer & Kreis 1978 

gracihs f. minor Patrick 1968 

gracihs var. schizonemoides Van Heurck Stoermer & Kreis 1978 

graciUima Ralfs Tempere & Peragallo 1911 

graciloides A. Mayer Stoermer & Kreis 1978 

gravistriata Patrick Patrick 1959 

gregaria Donkin Stoermer & Kreis 1978 

grimmei Krasske Stoermer & Kreis 1978 

guatemalensis Cleve & Grove Patrick & Reimer 1966 

guttata Grunow Patrick & Reimer 1966 

gysingensis Foged Stoermer & Kreis 1978 

habena Hohn & Hellerman Hohn & Hellerman 1963 

halophila (Grunow) Cleve Stoermer & Kreis 1978 

halophila var. minor Kolbe Patrick 1945 

halophila var. subcapitata 0strup Collins & Kalinsky 1977 

halophila f. tenuirostris Hustedt Camburn 1982 

hambergii Hustedt Stoermer & Kreis 1978 

harderi Hustedt Prescott & Dillard 1979 

hasta var. punctata Boyer Patrick & Reimer 1966 

hassiaca Krasske Stoermer & Kreis 1978 

hasta Pantocsek Stoermer & Kreis 1978 

hyalosirella Hustedt Hustedt 1962 

hebes Ralfs Cleve & MoUer 1878 

helensis Schulz Stoermer & Kreis 1978 

hemiptera Kutzing Stoermer & Kreis 1978 

hemiptera var. troiana Grunow Cleve & Moller 1879 

heroina A. Schmidt Patrick & Reimer 1966 

heufleri Grunow Stoermer & Kreis 1978 

heufleri var. leptocephala (Brebisson) Patrick Stoermer & Kreis 1978 

hexapla A. Schmidt Patrick & Reimer 1966 

hilseana Janisch Tempere & Peragallo 1912 

hitchcockii Ehrenberg Stoermer & Kreis 1978 

hudsonis Grunow Kalinsky 1983 

humerosa Brebisson Patrick & Reimer 1966 

humihs Donkin Boyer 1927b 

hungarica Grunow Stoermer & Kreis 1978 

hungarica var. capitata (Ehrenberg) Cleve Stoermer & Kreis 1978 

hungarica var. linearis 0strup Stoermer & Kreis 1978 

hustedtii Krasske Camburn 1982 

hustedtii f. obtusa (Hustedt) Hustedt Stoermer & Kreis 1978 

hustedtii f. philippina Skvortzow Hohn & Hellerman 1963 

hyalinula De Toni Kalinsky 1983 

icostauron (Ehrenberg) O'Meara Van Heurck & Grunow 1881-1885 (#138) 

illinoensis Ehrenberg 1856 

imbellis Hohn & Hellerman Hohn & Hellerman 1963 

imbricata Bock Stoermer & Kreis 1978 

ingrata Krasske Prescott & Dillard 1979 

importuna Hustedt Stoermer et al. 1999 

incerta Grunow Stoermer & Kreis 1978 

incomitatus Hohn & Hellerman Patrick & Reimer 1966 



446 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 



Publication 



Name 

Navicula incomposita Hagelstein Kalinsky 

Navicula incompta van incurva Reimer Reimer 

Navicula indemnis Hohn & Hellerman Patrick & Reimer 

Navicula indianensis Reimer Patrick & Reimer 

Navicula indifferens Hustedt Hohn & Hellerman 

Navicula inflata Donkin Stoermer & Kreis 

Navicula inflexa (Gregory) Ralfs Stoermer & Kreis 

Navicula infrenis Hohn & Hellerman Hohn & Hellerman 

Navicula ingenua Hustedt Stoermer & Kreis 

Navicula ingrata Krasske Stoermer & Kreis 

Navicula insignita Hustedt Stoermer & Kreis 

Navicula insociabilis Krasske Stoermer & Kreis 

Navicula insociabilis var. dissipatoides Hustedt Stoermer & Kreis 

Navicula instabilis A. Schmidt Tempere & Peragallo 

Navicula Integra (W. Smith) Ralfs Stoermer & Kreis 

Navicula interglacialis Hustedt Stoermer et al. 

Navicula intermedia Lagerstedt Elmore 

Navicula interrupta W. Smith Tempere & Peragallo 

Navicula interrupta var. stauroneiformis Tempere & Peragallo 

Navicula intracata Hustedt Stoermer & Kreis 

Navicula iridis Ehrenberg Stoermer & Kreis 

Navicula iridis var. affmis Ehrenberg Stoermer & Kreis 

Navicula iridis var. amphigomphus Ehrenberg Stoermer & Kreis 

Navicula iridis var. amphirhynchus (Ehrenberg) Cleve Stoermer & Kreis 

Navicula iridis var. ampliata Ehrenberg Tempere & Peragallo 

Navicula iridis var. firma W. Smith Stoermer & Kreis 

Navicula iridis var. maxima Tempere & Peragallo 

Navicula iridis var. producta W. Smith Stoermer & Kreis 

Navicula illinoensis Ehrenberg 

Navicula isocephala Ehrenberg Cleve & Moller 

Navicula jaagi Meister Stoermer & Kreis 

Navicula jaerenfeltii Hustedt Stoermer & Kreis 

Navicula jentzschii Grunow Stoermer & Kreis 

Navicula johnsonii O'Meara Aubert 

Navicula karsia Hohn Patrick & Reimer 

Navicula keeleyi Patrick Camburn 

Navicula kincaidii Sovereign Patrick & Reimer 

Navicula kisber Hohn & Hellerman Patrick & Reimer 

Navicula kotschyi Grunow Stoermer & Kreis 

Navicula krasskei Hustedt Stoermer & Kreis 

Navicula lacunarum Grunow Stoermer & Kreis 

Navicula lacustris Gregory Stoermer & Kreis 

Navicula laevissima Kiitzing Stoermer & Kreis 

Navicula laevissima f. fusticulus (0strup) Camburn Camburn et al. 

Navicula ladogensis Cleve Stoermer & Kreis 

Navicula lagerheimii Cleve Stoermer 

Navicula lagerheimii var. intermedia Hustedt Hohn & Hellerman 

Navicula lagerstedtii Cleve Patrick & Reimer 

Navicula lalia Hohn & Hellerman Hohn & Hellerman 

Navicula lanceolata (Agardh) Kutzing Stoermer & Kreis 

Navicula lanceolata \ar. cymbula (Donkin) Cleve Stoermer & Kreis 

Navicula lanceolata f. minuta Rabenhorst Patrick & Reimer 

Navicula lanceolata var. tenuirostris Stoermer et al. 

Navicula lapidosa Krasske Collins & Kalinsky 

Navicula lata (Brebisson) Kiitzing Elmore 

Navicula latelongitudinalis Patrick Patrick 

Navicula latens Krasske Stoermer & Kreis 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 447 



Name Publication 

Navicula lateropunctata Wallace Stoermer & Kreis 1978 

Navicula laterostrata Hustedt Stoermer & Kreis 1978 

Navicula latevittata Cleve Tempere & Peragallo 1908 

Navicula latissima Gregory Patrick & Reimer 1966 

Navicula legumen Ehrenberg Elmore 1922 

Navicula lenzii Hustedt Stoermer et al. 1999 

Navicula leptoceros Ehrenberg 1856 

Navicula leptogongyla Ehrenberg Kalinsky 1983 

Navicula leptorhynchus Ehrenberg Patrick & Reimer 1966 

Navicula leptosigma Ehrenberg Patrick & Reimer 1966 

Navicula leptostriata Jorgensen Dixit & Smol 1995 

Navicula leptotermia Ehrenberg 1 856 

Navicula levanderi Hustedt Stoermer & Kreis 1978 

Navicula libumica Grunow Elmore 1922 

Navicula limosa Kutzing Stoermer & Kreis 1978 

Navicula limosa var. gibberula (Kutzing) Grunow Stoermer & Kreis 1978 

Navicula limosa var. subinflata Grunow Stoermer & Kreis 1978 

Navicula limosa var. undulata Grunow Stoermer & Kreis 1978 

Navicula Hneolata Ehrenberg 1856 

Navicula lirata Ehrenberg Patrick & Reimer 1966 

Navicula litos Hohn & Hellerman Camburn 1982 

Navicula longa Ralfs Stoermer & Kreis 1978 

Navicula longicephala Hustedt Johansen et al 1983 

Navicula longirostris Hustedt Stoermer & Kreis 1978 

Navicula lucidula Grunow Patrick & Reimer 1966 

Navicula ludloviana A. Schmidt Boyer 1927b 

Navicula lundstroemii Cleve Stoermer & Kreis 1978 

Navicula luzonensis Hustedt Stoermer & Kreis 1978 

Navicula lyra Ehrenberg Hohn 195 1 

Navicula lyra var. elliptica A. Schmidt Patrick & Reimer 1966 

Navicula macilenta Cleve Tempere & Peragallo 1909 

Navicula maculata var. lanceolata Heiden Patrick & Reimer 1966 

Navicula maculata var. orbiculata Patrick Patrick & Reimer 1966 

Navicula major Kiitzing Stoermer & Kreis 1978 

Navicula major var. asymetrica Tempere & Peragallo 1909 

Navicula major var. dilatata M. Peragallo Patrick & Reimer 1966 

Navicula major var. maxima Patrick & Reimer 1966 

Navicula mandumensis Jorgensen Camburn & Charles 2000 

Navicula margaritaceae Hustedt Johansen et al. 1983 

Navicula maxima Gregory Stoermer & Kreis 1978 

Navicula meandrinoides Hustedt Hustedt 1930 

Navicula mediocris Krasske Stoermer & Kreis 1978 

Navicula mediocris var. intermedia Reimer Reimer 1966 

Navicula mediacomplexa Hohn & Hellerman Patrick & Reimer 1966 

Navicula mediahelos Hohn & Hellerman Hohn & Hellerman 1963 

Navicula medioconvexa Hustedt Potapova & Charles 2002 

Navicula megaloptera Ehrenberg Tempere & Peragallo 1908 

Navicula menisculoides Hustedt Stoermer et al. 1999 

Navicula menisculus Schumann Stoermer & Kreis 1978 

Navicula menisculus var. krenneri A. Cleve Stoermer et al. 1999 

Navicula menisculus f. linearis Reimer Reimer 1970 

Navicula meniscula var. muralis (Grunow) Lange-Bertalot Yearsley et al. 1992 

Navicula menisculus var. obtusa Hustedt Stoermer & Kreis 1978 

Navicula menisculus var. upsaliensis (Grunow) Grunow Camburn 1982 

Navicula meniscus Schumann Stoermer et al. 1999 

Navicula mesogongyla Ehrenberg Kalinsky 1983 

Navicula mesogongyla var. interrupta Cleve Tempere & Peragallo 1908 



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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 



Name 

Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu' 
Navicu 
Navicu^ 
Navicu' 
Navicu 
Navicu: 
Navicu^ 
Navicu^ 
Navicu! 
Navicu! 
Navicu! 
Navicu! 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu! 
Navicu 
Navicu! 
Navicu 
Navicu 
Navicu! 
Navicu! 
Navicu! 
Navicu 
Navicu 
Navicu 
Navicu! 
Navicu 
Navicu 
Navicu 
Navicu! 
Navicu 
Navicu 
Navicu 
Navicu! 
Navicu 
Navicu 
Navicu! 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 



Publication 



a mesolepta Ehrenberg Stoermer & Kreis 

a mesolepta var. stauroneiformis Grunow Tempere & Peragallo 

a mesostyla Ehrenberg Tempere & Peragallo 

a mica Hohn & Hellerman Hohn & Hellerman 

a micropupula Cholnoky Stoermer & Kreis 

a microstauron var. stauroneiformis Tempere & Peragallo 

a migma Hohn & Hellerman Hohn & Hellerman 

a minima Grunow Stoermer & Kreis 

a minima var. atomoides (Grunow) Cleve Patrick 

a minima var. okamurae Skvortzow Stoermer & Kreis 

a minima var. pseudofossilis (Krasske) Reimer Reimer 

a minnewaukonensis Elmore Stoermer & !Kreis 

a minthe Hohn & Hellerman Hohn & Hellerman 

a minuscula Grunow Stoermer & Kreis 

a minuscula f. linearis Reimer Reimer 

a minuscula var. alpestris Hustedt Stoermer & Kreis 

a minusculoides Hustedt Stoermer & Kreis 

a minuta Patrick & Reimer 

a mobiliensis Boyer Collins & Kalinsky 

a mobiliensis var. minor Patrick Patrick 



a modica Hustedt Cambum & Charles 2000 



a molestiformis Hustedt Grimes & Rushforth 



a monmouthiana-stodderi Yermeloff Boyer 1927b 



a monoculata Hustedt Stoermer & Kreis 

a montana Moghadam Prescott & Dillard 

a moumei Patrick Patrick 

a mucronata Elmore Elmore 

a multigramme Hohn & Hellerman Hohn & Hellerman 

a muraliformis Hustedt Stoermer & Kreis 

a muralis Grunow Patrick 

a murrayi West & West Rushforth & Merkley 

a muscerda Hohn Patrick & Reimer 

a mutica Kiitzing Stoermer & Kreis 

a mutica var. binodis Hustedt Cambum 

a mutica var. cohnii (Hilse) Grunow Stoermer & Kreis 

a mutica var. gibbosa IVIcCall Collins & Kalinsky 

a mutica var. goeppertiana (Bleisch) Grunow Collins & Kalinsky 

a mutica f. intermedia Hustedt Reimer 

a mutica var. nivalis (Ehrenberg) Hustedt Stoermer & Kreis 

a mutica var. stigma Patrick Patrick 

a mutica var. tropica Hustedt Collins & Kalinsky 

a mutica var. tropica f. rostrata Krasske Lowe & Kociolek 

a mutica var. undulata (Hilse) Grunow Stoermer & Kreis 

a mutica var. ventricosa Kutzing Hohn 

a mutica f. lanceolata Frenguelli Cambum 

a muticoides Hustedt Stoermer & Kreis 

a muticopsis Van Heurck Stoermer & Kreis 

a narinosa Hohn Patrick & Reimer 



a naumannii Hustedt Cambum & Charles 2000 

a nemoris Hohn & Hellerman Hohn & Hellerman 1963 

a neoventricosa Hustedt Stoermer & Kreis 1978 

a nigrii de Notaris Collins & Kalinsky 1977 

a nimbus Hohn & Hellerman Hohn & Hellerman 1963 

a nivalis Ehrenberg Collins & Kalinsky 1977 

a nobilis (Ehrenberg) Kutzing Stoermer & Kreis 1978 

a nobilis var. dactylus (Ehrenberg) Van Heurck Myers 1898b 

a nodosa Ehrenberg Elmore 1922 



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KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 449 



Name Publication 



Navicula nodosa f. curta Rabenhorst Tempere & Peragallo 

Navicula nodulosa Kiitzing Cleve & Moller 

Navicula nolens Simonsen Kalinsky 

Navicula notha Wallace Stoermer & Kreis 

Navicula nugalis Hohn & Hellerman Hohn & Hellerman 

Navicula nyassensis O. Mliller Stoermer & Kreis 

Navicula nyassensis var. capitata O. Miiller Hohn & Hellerman 

NaNicula nyassensis f. minor O. Miiller Stoermer & Kreis 

Navicula obdurata Hohn & Hellerman Stoermer & Kreis 

Navicula oblonga (Kutzing) Kutzing Stoermer & Kreis 

Navicula oblongata Kutzing Patrick & Reimer 

Navicula oblongiformis Hustedt Hustedt 

Navicula oblongum var. subcapitata Pantocsek Stoermer & Kreis 

Navicula obsidialis Hustedt Reimer 

Navicula obsoleta Hustedt Reimer 

Navicula obtusa Ehrenberg 

Navicula obtuseprotracta Hustedt 

Navicula ocallii Hohn Patrick & Reimer 

Navicula oculata Krasske Patrick & Reimer 

Navicula odiosa Wallace Stoermer & Kreis 

Navicula ohiensis Ehrenberg Patrick & Reimer 



Navicula okadae (Skvortzow) Nagumo & Kobayasi Cambum & Charles 2000 



Navicula omissa Hustedt Collins & Kalinsky 

Navicula opportuna Hustedt Stoermer et al. 

Navicula oppugnata Hustedt Stoermer & Kreis 

Navicula orangiana Patrick Patrick 

Navicula orbiculata Patrick Patrick 

Navicula ordinaria Hustedt Stoermer & Kreis 

Navicula oregonica Ehrenberg 

Navicula oxigua (Gregory) Miiller Patrick & Reimer 

Navicula paanaensis A. Cleve Stoermer & Kreis 

Navicula paca Hohn & Hellerman Stoermer & Kreis 

Navicula pachyptera Ehrenberg Tempere & Peragallo 

Navicula palpebralis Brebisson Elmore 

Navicula paludosa Hustedt Stoermer & Kreis 

Navicula paludosa f. rhomboidea Reimer Reimer 

Navicula paludosa var. rhomboidea Hustedt Collins & Kalinsky 

Navicula pampeana Frenguelli Stoermer 

Navicula parablis Hohn & Hellerman Hohn & Hellerman 

Navicula paramutica Bock Rushforth & Merkley 

Navicula paramutica var. binodis Bock Lawson & Rushforth 



Navicula parasubtilissima Kobayasi & Nagumo Cambum & Charles 2000 

Navicula paratunkae Peterson Cambum 1982 

Navicula parodia Hohn Patrick & Reimer 1966 

Navicula parva Ralfs Stoermer & Kreis 1978 

Navicula parva (Ehrenberg) Elmore Elmore 1922 

Navicula parva (Meneghin) Cleve Kaczmarska & Rushforth 1983 

Navicula paucivisitata Patrick Patrick 1959 

Navicula paulensis Gmnow Tempere & Peragallo 1913 

Navicula pavillardi Hustedt Kalinsky 1983 

Navicula pelliculosa Hilse Stoermer & Kreis 1978 

Navicula pennata A. Schmidt Patrick & Reimer 1966 

Navicula pennsylvanica Patrick Patrick & Reimer 1966 

Navicula peratomus Hustedt Stoermer & Kreis 1978 

Navicula peregrina (Ehrenberg) Kutzing Stoermer & Kreis 1978 

Navicula peregrina var. truncata M. Peragallo in Tempere & Peragallo Tempere & Peragallo 1908 

Navicula peripunctata J. Brun Tempere & Peragallo 1908 



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Publication 



Name 

Navicula permitis Hustedt Stoermer & Kreis 

Navicula perpusilla Grunow Stoermer & Kreis 

Navicula perpusilla var. distans Cleve-Euler Patrick & Reimer 

Navicula perrotettii (Grunow) Cleve Stoermer & Kreis 

Navicula perrotettii var. enervis Hustedt Stoermer et al. 

Navicula perventralis Hustedt Patrick & Reimer 

Navicula peticolasii M. Peragallo Patrick & Reimer 

Navicula phyllodes Ehrenberg 

Navicula placenta Ehrenberg Stoermer & Kreis 

Navicula placentula (Ehrenberg) Kiitzing Stoermer & Kieis 

Navicula placentula var. jenisseyensis (Grunow) Meister Hohn 

Navicula placentula var. latiuscula (Grunow) Meister Drum 

Navicula placentula var. maculata Hustedt Stoermer et al. 

Navicula placentula var. rostrata A. Mayer Stoermer & Kreis 

Navicula placentula f. rostrata A. Mayer Cambum 

Navicula platalea Ehrenberg Patrick & Reimer 

Navicula platycephala O. Muller Stoermer & Kreis 

Navicula platysoma Ehrenberg Stoermer & Kreis 

Navicula platysoma var. pantocsekii Wislough & Kolbe Stoermer & Kreis 

Navicula platyventris Meister Kalinsky 

Navicula pletura Hohn Patrick & Reimer 

Navicula poconoensis Patrick Patrick 

Navicula pollis Van Heurck & Grunow 1882-1885 

Navicula polyonca Brebissona Aubert 

Navicula polystricta var. circumstricta Grunow Collins & Kalinsky 

Navicula porifera var. oppotuna (Hustedt) Lange-Bertalot Stoermer et al. 

Navicula portomonttana Cleve Bateman & Rushforth 

Navicula potzgeri Reimer Stoermer & Kreis 

Navicula potgeri var. quadripunctata Reimer Patrick & Reimer 

Navicula praeterita Hustedt Stoermer et al. 

Navicula pragma Hohn & Hellerman Hohn & Hellerman 

Navicula producta W. Smith Stoermer & Kreis 

Navicula protracta (Grunow) Cleve Stoermer & Kreis 

Navicula protracta var. elliptica Gallik Stoermer & Kreis 

Navicula protracta f. subcapitata (Wils. & Por.) Hustedt Stoermer & Kreis 

Navicula pseuanglica Lange-Bertalot Stoermer et al. 

Navicula pseudoarvensis Hustedt Cambum et al. 

Navicula pseudoatomus Lund Loescher 

Navicula pseudobacillum Grunow Stoermer & Kreis 

Navicula pseudocanalis Patrick & Roberts 

Navicula pseudoclementis Hustedt Stoermer & Kreis 

Navicula pseudocrassirostris Hustedt Kaczmarska & Rushforth 

Navicula pseudoexillissima Hustedt Fee 

Navicula pseudofrickia Patrick Patrick & Reimer 

Navicula pseudolanceolata Lange-Bertalot Siver et al. 

Navicula pseudomuralis Hustedt Stoermer et al. 

Navicula pseudopelliculosa Manguin Hohn & Hellerman 

Navicula pseudoreinhardtii Patrick Stoermer & Kreis 

Navicula pseudoscutiformis Hustedt Stoermer & Kreis 

Navicula pseudosilicula Hustedt Bateman & Rushforth 

Navicula pseudosilicula var. olympica Sovereign Patrick & Reimer 

Navicula pseudosubtilissima Manguin Cambum & Charles 

Navicula pseudotuscula Hustedt Grimes & Rushforth 

Navicula pseudoventralis Hustedt Stoermer & Kreis 

Navicula punctata Donkin Stoermer & Kreis 

Navicula pupula Kiitzing Stoermer & Kreis 

Navicula pupula var. aquaeductae (Krasske) Hustedt Stoermer & Kreis 



(#5 



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44) 
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KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 451 



Name Publication 

Na\icula pupula var. bacillarioides Grunow Stoermer & Kreis 1978 

Navicula pupula var. capitata Hustedt Stoenner & Kreis 1978 

Navicula pupula var. elliptica Hustedt Stoermer & Kreis 1978 

Navicula pupula var. lineare Tempere & Peragallo Tempere & Peragailo 1908 

Navicula pupula var. major O. Muller Patrick & Reimer 1966 

Navicula pupula var. minor Kiitzing Tempere & Peragallo 1908 

Navicula pupula f. minor Tempere & Peragallo 1909 

Navicula pupula f. minuta Tempere & Peragailo 1909 

Navicula pupula var. minuta Van Heurclc Clark & Rushforth 1977 

Navicula pupula f. minutula Cholnoky Hohn & Hellerman 1963 

Navicula pupula var. mutata (Krasske) Hustedt Stoermer & Kreis 1978 

Navicula pupula var. rectangularis (Gregory) Cleve Stoermer & Kreis 1978 

Navicula pupula var. rostrata Hustedt Stoermer & Kreis 1978 

Navicula pupula f. rostrata Hustedt Camburn 1982 

Navicula pusilla W. Smith Stoermer & Kreis 1978 

Navicula pusilla var. lanceolata (Grunow) Grunow Patrick & Reimer 1966 

Navicula pusio Cleve Prescott & Dillard 1979 

Navicula pygmaea Kiitzing Stoermer & Kreis 1978 

Navicula quadripartita Hustedt Stoermer & Kreis 1978 

Navicula rabenhorstii Ralfs Aubert 1895 

Navicula radians Patrick & Reimer 1966 

Navicula radiosa Kutzing Stoermer & Kreis 1978 

Navicula radiosa var. acuta Grunow Tempere & Peragallo 1908 

Navicula radiosa var. parva Wallace Stoermer & Kreis 1978 

Navicula radiosa var. subrostrata Cleve Patrick & Reimer 1966 

Navicula radiosa var. tenella (Brebisson) Grunow Stoermer & Kreis 1978 

Navicula radiosafallax Lange-Bertalot Stoermer et al. 1999 

Navicula rainierensis Sovereign Patrick & Reimer 1966 

Navicula rangoonensis (Grunow) Elmore Elmore 1922 

Navicula recava Hohn & Hellerman Hohn & Hellerman 1963 

Navicula recens Lange-Bertalot Potapova & Charles 2003 

Navicula recondita Torka Stoermer & Kreis 1978 

Navicula reichardtiana Lange-Bertalot Potapova & Charles 2003 

Navicula reinhardtii Grunow Stoermer & Kreis 1978 

Navicula reinhardtii var. elliptica Heribaud Stoermer & Kreis 1978 

Navicula retusa var. elongata Patrick & Reimer 1966 

Navicula rhodana Hohn & Hellerman Patrick & Reimer 1966 

Navicula rhomboides Grunow Stoermer & Kreis 1978 

Navicula rhomboides var. major Cleve & Moller 1879 

Navicula rhynchocephala Kutzing Stoermer & Kreis 1978 

Navicula rhynchocephala var. amphiceros (Kutzing) Grunow Stoermer & Kreis 1978 

Navicula rhynchocephala var. germainii (Wallace) Patrick Stoermer & Kreis 1978 

Navicula rhynchotella Lange-Bertalot Stoermer et al. 1999 

Navicula rivalis Hohn & Hellerman Hohn & Hellerman 1963 

Navicula rostellata Kutzing Stoermer & Kreis 1978 

Navicula rostrata Ehrenberg Elmore 1922 

Navicula rotaeana (Rabenhorst) Grunow Stoermer & Kreis 1978 

Navicula rotaeana var. excentrica Grunow Stoermer & Kreis 1978 

Navicula romnda Hustedt Stoermer & Kreis 1978 

Navicula rugula Hohn & Hellerman Hohn & Hellerman 1963 

Navicula rupestris Hantzsch Myers 1 898b 

Navicula ruttneri Hustedt Sovereign 1958 

Navicula sabiniana Patrick Stoermer & Kreis 1978 

Navicula sagitta Hohn & Hellerman Hohn & Hellerman 1963 

Navicula salinarum Grunow Stoermer & Kreis 1978 

Navicula salinarum var. capitata Schuiz Kaczmarska & Rushforth 1983 

Navicula salinarum var. intermedia (Grunow) Cleve Stoermer & Kreis 1978 



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Name 

Navicu 
Navicu 
Navicu 
Navicu: 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navica 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 
Navicu 



Publication 

a salinicola Hustedt Potapova & Charles 2003 

982 
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982 
856 

a schadei Krasske Cambum & Charles 2000 



a sanctaecrucis 0strup Cambum 

a saprophila Lange-Bertalot & Bonik Johansen et al. 

a saugerii Desm Stoermer & Kreis 

a savannahiana Patrick Cambum 

a scalpmm Ehrenberg 



a schmassmannii Hustedt Stoem:ier & Kreis 

a schoenfeldii Hustedt Stoemier & Kreis 

a schroteri Meister Hohn & Hellerman 

a schroeteri var. escambia Patrick Stoermer & Kreis 

a schumanniana Grunow Stoermer & Kreis 

a schweinfurthii A. Schmidt Aubert 

a sculpta Ehrenberg Stoermer & Kreis 

a scutelloides W. Smith Stoermer & Kreis 

a scutelloides var. minutissima Tempere & Peragallo 

a scutelloides var. mocarensis Gmnow Patrick & Reimer 

a scutiformis Gmnow Stoermer & Kreis 

a scutula Hohn Patrick & Reimer 

a scutum Schumann Patrick & Reimer 

a secreta Pantocsek Stoermer & Kreis 

a secura Patrick Stoermer & Kreis 

a secreta var. apiculata Patrick Cambum 



a semen (Ehrenberg) Donkin Boyer 1927b 



a semenoides Hustedt Stoermer & Kreis 

a seminuloides Hustedt Stoermer & Kreis 

a seminuloides var. sumatrana Hustedt Sovereign 

a seminulum Gmnow Stoermer & Kreis 

a seminulum var. hustedtii Patrick Cambum 

a seminulum var. intermedia Hustedt Stoermer & Kreis 

a septata Hustedt Rushforth & Merkley 

a septenaria L.W. Bailey Patrick & Reimer 

a serians Kutzing Stoermer & Kreis 

a serians var. follis Ehrenberg Cleve & Moller 

a serians var. minor Tempere & Peragallo 

a sigma Ehrenberg Stoermer & Kreis 

a silicula Ehrenberg Elmore 

a similis Krasske Stoermer & Kreis 

a simplex Krasske Cambum 

a Simula Patrick Patrick 

a singularis Schmidt Cleve & Moller 

a skabitschewskyi Sabelina Stoermer & Kreis 

a skalenastriata Hohn Patrick & Reimer 

a spirata Hustedt Collins & Kalinsky 

a splendicula Van Landingham Cambum 

a smithii Brebisson Stoermer & Kreis 

a smithii var. dilatata M. Peragallo in Tempere & Pergallo Tempere & Peragallo 

a sohrensis Krasske Bateman & Rushforth 



a sohrensis var. hassiaca (Krasske) Lange-Bertalot Cambum & Charles 2000 



a sorella Hohn & Hellerman Patrick & Reimer 

a sovereignae Hustedt Stoermer & Kreis 

a speciosa Hustedt Hustedt 

a sphaerophora Kutzing Stoermer & Kreis 

a spicula (Hickie) Cleve Kalinsky 

a splendicula Van Landingham Stoermer et al. 

a stankovici Hustedt Patrick & Reimer 

a staurifera Thomas Stoermer & Kreis 



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KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 453 



Name Publication 



Navicula stauroneiformis Elmore Patrick & Reimer 

Navicula stauroptera Ralfs Tempere & Peragallo 

Navicula stauroptera var. parva (Ehrenberg) Grunow Stoermer & Kreis 

Navicula stauroptera var. troiana Grunow Cleve & Moller 

Navicula stodderi Greenl Cleve 

Navicula stomatophora Grunow Tempere & Peragallo 

Navicula strenzkei Hustedt Grimes & Rushforth 

Navicula stroemii Hustedt Stoermer & Kreis 

Navicula stroesei (0strup) Cleve Stoermer & Kreis 

Navicula subacuta Ehrenberg Tempere & Peragallo 

Navicula subadnata Hustedt Lawson & Rushforth 

Navicula subarxensis Hustedt Dodd 

Navicula subatomoides Hustedt Patrick 

Navicula subbacillum Hustedt Collins & Kalinsky 

Navicula subcapitata (Gregory) Ralfs Elmore 

Navicula subcapitata var. stauroneiformis Tempere & Peragallo 

Navicula subclementis Hustedt Stoermer & Kreis 

Navicula subcontenta Hustedt Reimer 

Navicula subcostulata Hustedt Stoermer & Kreis 

Navicula subfasciata Patrick Patrick 

Navicula subgastriformis Hustedt Stoermer et al. 

Navicula subhalophila Hustedt Stoermer & Kreis 

Navicula subhamulata Grunow Stoermer & Kreis 

Navicula subhamulata var. undulata Hustedt Stoermer & Kreis 

Navicula subhexagona Hustedt Hustedt 

Navicula subinflata Grunow Patrick & Reimer 

Navicula subinflatoides Hustedt Czamecki et al. 

Navicula sunminuscula Manguin Hohn & Hellerman 

Navicula submitis Hustedt Stoermer et al. 

Navicula submolesta Hustedt Patrick & Reimer 

Navicula submuralis Hustedt Stoermer & Kreis 

Navicula subocculata Hustedt Stoermer & Kreis 

Navicula subrhynchocephala Hustedt Stoermer & Kreis 

Navicula subrotundata Hustedt Stoermer & Kreis 

Navicula subrotundata f. lanceolata Hustedt Stoermer et al. 

Navicula subseminulum Hustedt Stoermer & Kreis 

Navicula subsulcata Hustedt Stoermer & Kreis 

Navicula subsulcatoides Hustedt Kaczmarska & Rushforth 

Navicula subtilissima Cleve Stoermer & Kreis 

Navicula suchlandtii Hustedt Lowe & Kociolek 

Navicula swaniana Moghadam Prescott & Dillard 

Navicula symmetrica Patrick Stoermer & Kreis 

Navicula tabellaria Ehrenberg Stoermer & Kreis 

Navicula tabellaria var. stauroneiformis Aubert 

Navicula taedia Wallich Patrick & Reimer 

Navicula tanmla Hustedt Stoermer & Kreis 

Navicula tantula var. minima Grunow Collins & Kalinsky 

Navicula taraxa Hohn & Hellerman Kalinsky 

Navicula tecta Krasske Stoermer & Kreis 

Navicula tenella (Brebissonj Van Heurck Stoermer & Kreis 

Navicula tenelloides Hustedt Collins & Kalinsky 

Navicula tenera Hustedt Collins & Kalinsky 

Navicula tenuicephala Hustedt Dixit & Smol 

Navicula termes Ehrenberg Stoermer & Kreis 

Navicula terminata Hustedt Stoermer & Kreis 

Navicula terrestris Petersen Dodd 

Navicula terrestris var. relicta f. triundulata Lund Dodd 



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Publication 



Name 

Navicula texana Patrick Stoermer & Kreis 

Navicula thermes (Ehrenberg) A. Schmidt Elmore 

Navicula thienemannii Hustedt Reimer 

Navicula tracery Hohn & Hellerman Patrick & Reimer 

Navicula transversa A. Schmidt Tempere & Peragallo 

Navicula tridentula Krasske Stoermer & Kreis 

Navicula triduentula var. parallela Krasske Stoermer & Kreis 

Navicula trinodis Lewis Stoermer & Kreis 

Navicula tripunctata (O.F. Miiller) Bory Stoermer & Kreis 

Navicula tripunctata var. cuneata (Lauby) Stoermer & Yang Stoermer & Kreis 

Navicula tripunctata var. schizonemoides (Van Heurck) Patrick Stoermer & Kreis 

Navicula trivialis Lange-Bertalot Yearsely et al. 

Navicula trochus Schumann Stoermer & Kreis 

Navicula tulugakii Carter Rushforth & Merkley 

Navicula tumida W. Smith Stoermer & Kreis 

Navicula tuscula Ehrenberg Stoermer & Kreis 

Navicula tuscula var. angulata Hustedt Stoermer et al. 

Navicula tuscula var. rostrata (Hustedt) Hustedt Stoermer et al. 

Navicula tuscula f. minor Hustedt Stoermer & Kreis 

Navicula tuscula f. obtusa (Hustedt) Hustedt Stoermer & Kreis 

Navicula tuscula f. rostrata Hustedt Stoermer & Kreis 

Navicula typografica Hustedt Hustedt 

Navicula umbra Hohn & Hellerman Hohn & Hellerman 

Navicula utermohleii Hustedt Prescott & Dillard 

Navicula vanheurckii Patrick Stoermer & Kreis 

Navicula varians Gregory Stoermer & Kreis 

Navicula variostriata Krasske Stoermer & Kreis 

Navicula vaucheriae J.B. Petersen Stoermer et al. 

Navicula venerablis Hohn & Hellerman Hohn & Hellerman 

Navicula veneta Kutzing Johansen et al. 

Navicula ventosa Hustedt Stoermer & Kreis 

Navicula ventralis Krasske Stoermer & Kreis 

Navicula ventralis var. chilensis Krasske Hohn & Hellerman 

Navicula ventralis f. simplex Hustedt Stoermer & Kreis 

Navicula ventricosa Kutzing Tempere & Peragallo 

Navicula ventricosa var. undulata Tempere & Peragallo 

Navicula verecunda Hustedt Stoermer & Kreis 

Navicula vetita Krasske Dodd 

Navicula viridis (Nitzsch) Ehrenberg Stoermer & Kreis 

Navicula viridis var. fallax Cleve Tempere & Peragallo 

Navicula viridis var. commutata Grunow Tempere & Peragallo 

Navicula viridula (Kutzing) Ehrenberg Stoermer & Kreis 

Navicula viridula var. argunensis Skvortzow Collins & Kalinsky 

Navicula viridula var. avenacea (Brebisson) Van Heurck Stoermer & Kreis 

Navicula viridula var. germainii (J.H. Wallace) Lange-Bertalot Lange-Bertalot 

Navicula viridula var. linearis Hustedt Stoermer & Kreis 

Navicula viridula var. rostellata (Kutzing) Cleve Stoermer & Kreis 

Navicula vitabunda Hustedt Stoermer & Kreis 

Navicula vitabunda var. montana Moghadam Prescott & Dillard 

Navicula vitiosa Schimanski Cambum & Charles 

Navicula vulpina Kutzing Stoermer & Kreis 

Navicula vulpina var. avenacea (Van Heurck) Patrick Stoermer et al. 

Navicula walked Sovereign Patrick & Reimer 

Navicula wallacei Reimer Cambum 

Navicula wardii Patrick Patrick 

Navicula wittrockii (Lagerstedt) Tempere & M. Peragallo Stoermer & Kreis 

Navicula wulffi Boye Peterson Hohn & Hellerman 



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KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 455 



Name Publication 

Navicula yarrensis Gninow Kalinsky 1983 

Navicula yorkensis Cambum Camburn 1982 

Navicula zanoni Hustedt Stoermer & Kreis 1978 

Neidium affine (Ehrenberg) Pfitzer Stoermer & Kreis 1978 

Neidium affine var. amphirhynchus (Ehrenberg) Cleve Stoermer & Kreis 1978 

Neidium affine var. bonsaensis Foged Lowe 1972-1973 

Neidium affine var. capitata Molder Stoermer et al. 1999 

Neidium affine var. ceylonicum (Skvortzow) Reimer Stoermer & Kreis 1978 

Neidium affine var. genuinum f. maxima Cleve Boyer 1927b 

Neidium affine var. hankense (Skvortzow) Reimer Stoermer & Kreis 1978 

Neidium affine var. humerus Reimer Stoermer & Kreis 1978 

Neidium affine var. longiceps (Gregory) Cleve Camburn 1982 

Neidium affine var. quadripunctatum (Hustedt) Hamilton Camburn & Charles 2000 

Neidium affine var. tenuirostris A. Mayer Stoermer 1963 

Neidium affine var. undulatum (Grunow) Cleve Stoermer & Kreis 1978 

Neidium alpinum Hustedt Patrick & Reimer 1966 

Neidium alpinum var. quadripunctatum (Hustedt) Hamilton Hamilton et al. 1990 

Neidium amphigomphus (Ehrenberg) Pfitzer Stoermer & Kreis 1978 

Neidium amphirhynchus (Ehrenberg) Boyer Boyer 1927b 

Neidium ampliatum (Ehrenberg) Krammer Stoermer et al. 1999 

Neidium apiculatum Reimer Stoermer & Kreis 1978 

Neidium apiculatum var. constrictum Reimer Patrick & Reimer 1966 

Neidium binode (Ehrenberg) Hustedt Stoermer & PCreis 1978 

Neidium bisulcatum (Lagestedt) Cleve Stoermer & Kreis 1978 

Neidium bisulcatum var. baicalense (Skvortzow & Meyer) Reimer Stoermer & Kreis 1978 

Neidium bisulcatum f. lineare (0strup) Cleve-Euler Bateman & Rushforth 1984 

Neidium bisulcatum var. nipponicum Skvortzow Patrick & Reimer 1966 

Neidium bisulcatum var. subundulatum (Grunow) Reimer Camburn 1982 

Neidium bisulcatum f. undulata (O. Miiller) Hustedt Bateman & Rushforth 1984 

Neidium bisulcatum var. undulatum O. Muller Patrick 1945 

Neidium boyeri Reimer Patrick & Reimer 1966 

Neidium calvum 0strup Stoermer & Kreis 1978 

Neidium cape-codii Siver & Hamilton in Siver et al Siver et al. 2003 

Neidium decens (Pantocsek) Stoermer Stoermer 1963 

Neidium densestriamm (0strup) Krammer Hamilton et al. 1992 

Neidium dilatamm (Ehrenberg) Cleve Boyer 1927b 

Neidium distincte-punctatum Hustedt Stoermer & Kreis 1978 

Neidium dubium (Ehrenberg) Cleve Stoermer & Kreis 1978 

Neidium dubium f. constrictum Hustedt Stoermer & Kreis 1978 

Neidium floridanum Reimer Patrick & Reimer 1966 

Neidium gracile f. aequale Hustedt Patrick & Reimer 1966 

Neidium hankensis Skvortzow Stoermer 1963 

Neidium hankensis var. elongata Skvortzow Stoermer 1963 

Neidium hercynicum A. Mayer Camburn 1982 

Neidium hercynicum f. subrostratum Wallace Camburn 1982 

Neidium herrmannii Hustedt Patrick & Reimer 1966 

Neidium hitchcockii (Ehrenberg) Cleve Stoermer & Kreis 1978 

Neidium hitchcockii f. teres Sovereign Patrick & Reimer 1966 

Neidium holstii (Cleve) Krammer Cambum & Charles 2000 

Neidium inconstans Sovereign Patrick & Reimer 1966 

Neidium iridis (Ehrenberg) Cleve Stoermer & Kreis 1978 

Neidium iridis var. amphigomphus (Ehrenberg) A. Mayer Stoermer & Kreis 1978 

Neidium iridis var. ampliatum (Ehrenberg) Cleve Stoermer & Kreis 1978 

Neidium iridis var. conspicua A. Mayer Patrick & Reimer 1966 

Neidium iridis var. intercedens A. Mayer Schmidt & Fee 1967 

Neidium iridis var. subundulatum (Cleve-Euler) Reimer Patrick & Reimer 1966 

Neidium iridis var. vemalis Reichelt Stoermer & Kreis 1978 



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Name Publication 

Neidium knuthii var. heilprinensis Foged Patrick & Reimer 1966 

Neidium kozolwi Mereschkowsky Stoermer & Kreis 1978 

Neidium kozolwi var. baicalensis f. robusta Stoermer Stoermer & Kreis 1978 

Neidium kozlowii var. parvum Mereschkowsky Patrick & Reimer 1966 

Neidium kozlowi var. undulata Stoermer Stoermer 1963 

Neidium ladogense (Cleve) Stoermer & Yang Stoermer & Kreis 1978 

Neidium ladogense var. densestriatum (0strup) Foged Cambum 1982 

Neidium ladogense var. densestriatum f. peribryum Lowe & Kociolek Lowe & Kociolek 1984 

Neidium levanderi (Hustedt) Lange-Bertalot & Metzeltin Stoermer et al. 1999 

Neidium magellanicum Cleve Prescott & Dillard 1979 

Neidium maximum (Cleve) Meister Patrick & Reimer 1966 

Neidium mirum Krasske Stoermer et al. 1999 

Neidium munckii Foged Lowe 1972-1973 

Neidium productum (W. Smith) Cleve Stoermer & Kreis 1978 

Neidium rudimentarum Reimer Patrick & Reimer 1966 

Neidium sacoense Reimer Stoermer & Kreis 1978 

Neidium temperei Reimer Stoermer & Kreis 1978 

Neidium tenuissimum Hustedt Johansen et al. 1983 

Neidium tumescens (Grunow) Cleve Boyer 1927b 



Nitzsch 
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Nitzsch 
Nitzsch 
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Nitzsch 
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a abbreviata Hustedt Collins & Kalinsky 1977 

a abridia Cambum Cambum 1 982 

a accedens Hustedt Hohn & Hellerman 1963 

a accomodata Hustedt Stoermer & Kreis 1978 

a acicuiarioides Hustedt Stoermer & Kreis 1978 

a acicularis (Kutzing) W. Smith Stoermer & Kreis 1978 

a acicularis var. adelos Hohn & Hellerman Hohn & Hellerman 1963 

a acicularis closterioides Grunow Czamecki & Blinn 1978 

a acida Cambum Johansen et al. 2004 

a acidoclinata Lange-Bertalot Stoermer et al. 1999 

a actinastroides (Lemmerman) Van Goor Stoermer & Kreis 1978 

a acula Hantzsch Dodd 1987 

a acuminata ( W. Smith) Grunow Stoermer & Kreis 1978 

a acuta Hantzsch Stoermer & Kreis 1978 

a acutiuscula Grunow Tempere & Peragallo 1909 

a adamata Hustedt Hansmann 1973 

a adapta Hustedt Collins & Kalinsky 1977 

a admissoides Cholnoky Collins & Kalinsky 1977 

a aerophila Hustedt Potapova & Charles 2002 

a agnewii Cholnoky Stoermer & Kreis 1978 

a agnita Hustedt Collins & Kalinsky 1977 

a amphibioides Hustedt Cambum et al. 1978 

a amphibia Grunow Stoermer & Kreis 1978 

a amphibia var. fossilis Gmnow Stoermer & Kreis 1978 

a amphibia var. frauenfeldii Gmnow Stoermer et al. 1999 

a amphicephala Gmnow Stoermer & Kreis 1978 

a amphioxoides Hustedt Stoermer et al. 1999 

a amphiox>'s (Ehrenberg) W. Smith Stoermer & Kreis 1978 

a amplectens Hustedt Cambum 1982 

a annularis var. affmis Gmnow Benson & Rushforth 1975 

a angustata (W. Smith) Gmnow Stoermer & Kreis 1978 

a angustata var. acuta Gmnow Stoermer & Kreis 1978 

a angustulata Lange-Bertalot Stoermer et al. 1999 

a apiculata (Gregor)) Gmnow Stoermer & Kreis 1978 

a archibaldii Lange-Bertalot Potapova & Charles 2002 

a aurariae Cholnoky Potapova & Charles 2002 

a asymbasia Hohn Hohn 1961 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 457 



Name Publication 

Nitzschia bacata Hustedt Stoermer & Kreis 1978 

Nitzschia bacata f. linearis Hustedt Stoermer & Kreis 1978 

Nitzschia bacillariaeformis Collins & Kalinsky 1977 

Nitzschia bacillariaeformis var. producta Cholnoky Collins & Kalinsky 1977 

Nitzschia balcanica Hustedt Johansen et al. 1983 

Nitzschia bella Sovereign Sovereign 1963 

Nitzschia bergii A. Cleve Czarnecki et al. 1981 

Nitzschia biacrula Hohn & Hellerman Camburn 1982 

Nitzschia bicrena Hohn & Hellerman Hohn & Hellerman 1963 

Nitzschia bilobata var. ambigua Manguin Kalinsky 1983 

Nitzschia bita Hohn & Hellerman Hohn & Hellerman 1963 

Nitzschia brebissonii W. Smith Boyer 1927b 

Nitzschia bremensis Hustedt Collins & Kalinsky 1977 

Nitzschia brevirostris Hustedt Collins & Kalinsky 1977 

Nitzschia brevissima (Grunow) Kalinsky Camburn 1982 

Nitzschia bryophila Hustedt Reimer 1990 

Nitzschia bulnheimiana (Rabenhorst) H.L. Smith Stoermer & Kreis 1978 

Nitzschia bulnheimiana var. capitata Reimer Reimer 1966 

Nitzschia caledonensis Schoeman Dodd 1987 

Nitzschia calida Grunow Camburn 1982 

Nitzschia capitellata Hustedt Stoermer & Kreis 1978 

Nitzschia chasei Cholnoky Collins & Kalinsky 1977 

Nitzschia circumsuta (Bailey) Grunow Stoermer et al. 1999 

Nitzschia clausii Hantzsch Stoermer & Kreis 1978 

Nitzschia closterium (Ehrenberg) W. Smith Stoermer & Kreis 1978 

Nitzschia coarctata Grunow Camburn 1982 

Nitzschia columbiana Sovereign Sovereign 1958 

Nitzschia communis Rabenhorst Stoermer & Kreis 1978 

Nitzschia conmiunis var. abbreviata Grunow Collins & Kalinsky 1977 

Nitzschia communis var. hyalina Lund Reimer 1970 

Nitzschia communis var. obtusa Grunow Reimer 1961 

Nitzschia commutata Grunow Stoermer & Kreis 1978 

Nitzschia compressa (Bailey) Boyer Collins & Kalinsky 1977 

Nitzschia confmis Hustedt Stoermer & Kreis 1978 

Nitzschia congolinensis Hustedt Wujek & Rupp 1980 

Nitzschia congolensis var. mooreae Reimer Reimer 1982 

Nitzschia constricta (Kutzing) Ralfs Camburn 1982 

Nitzschia constricta var. subconstricta Grunow Camburn 1982 

Nitzschia debilis Amott ex Grunow Camburn 1982 

Nitzschia debilis H.L. Smith H.L. Smith 1876-1888 (#690) 

Nitzschia delicatissima Cleve Stoermer & Kreis 1978 

Nitzschia denticula Grunow Stoermer & Kreis 1978 

Nitzschia denticula var. aberrans Fusey Reimer 1961 

Nitzschia desertorum Hustedt Collins & Kalinsky 1977 

Nitzschia diserta Hustedt Stoermer & Kreis 1978 

Nitzschia dissipata fKutzing) Grunow Stoermer & Kreis 1978 

Nitzschia dissipata var. bomeensis Hustedt Stoermer & Kreis 1978 

Nitzschia dissipata var. media Grunow Stoermer & Kreis 1978 

Nitzschia dissipata f. undulata Sovereign Camburn 1982 

Nitzschia draveillensis Coste & Ricard Stoermer et al. 1999 

Nitzschia dubia W. Smith Stoermer & Kreis 1978 

Nitzschia elegans Hustedt Stoermer & Kreis 1978 

Nitzschia elegantula Grunow Camburn 1982 

Nitzschia elliptica Hustedt Collins & Kalinsky 1977 

Nitzschia epiphytica Hustedt Stoermer & Kreis 1978 

Nitzschia epiphyticoides Hustedt Stoermer & Kreis 1978 

Nitzschia epithemioides Grunow Rushforth & Squires 1985 



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Name Publication 

Nitzschia etoshensis Cholnoky Collins & Kalinsky 

Nitzschia exilis Sovereign Sovereign 

Nitzschia fasciculata Grunow Stoermer & Kreis 

Nitzschia filiformis (W. Smith) Schutt Stoermer & Kreis 

Nitzschia filiformis var. conferta (P.G. Richter) Lange-Bertalot Stoermer et al. 

Nitzschia flexa Schumann Stoermer & Kreis 

Nitzschia fluminenensis Grunow Stoermer & Kreis 

Nitzschia fonticola Grunow Stoermer & Kreis 

Nitzschia fonticola var. capitata A. Cleve Stoermer & Kreis 

Nitzschia fonticula var. pelagica Hustedt Stoermer & Kreis 

Nitzschia fonticola var. romana (Grunow) Cleve-Euler Whitford & Schumacher 

Nitzschia fonticoloides Sovereign Stoermer & Kreis 

Nitzschia frequens Hustedt Collins & Kalinsky 

Nitzschia frustulum (Kiitzing) Grunow Stoermer & Kreis 

Nitzschia frustulum var. indica Patrick 

Nitzschia frustulum var. minutula Stoermer et al. 

Nitzschia frustulum var. perminuta Grunow Stoermer & Kreis 

Nitzschia frustulum var. perpusilla (Rabenhorst) Grunow Stoermer & Kreis 

Nitzschia frustulum var. subsalina Hustedt Stoermer & Kreis 

Nitzschia frustulum var. subserians Patrick & Roberts 

Nitzschia frustulum var. tenella Grunow Stoermer & Kreis 

Nitzschia gandersheimiensis Krasske Stoermer & Kreis 

Nitzschia geitleri Hustedt Dodd 

Nitzschia gessneri Hustedt Stoermer et al. 

Nitzschia graciliformis Lange-Bertalot & Simonsen Stoermer et al. 

Nitzschia graciloides Hustedt Stoermer et al. 

Nitzschia gracilis Hantzsch Stoermer & Kreis 

Nitzschia gracilis var. minor Patrick 

Nitzschia gracilis var. schizonemoides Van Heurck Starrett & Patrick 

Nitzschia hantzschiana Rabenhorst Stoermer & Kreis 

Nitzschia hantzschiana f. subserians Grunow Patrick 

Nitzschia hennedyi W. Smith Whitford & Schumacher 

Nitzschia heutleuriana Grunow Stoermer & Kreis 

Nitzschia hiemalis Hustedt Reimer 

Nitzschia hollerupensis Foged Stoermer et al. 

Nitzschia holsatica Hustedt Stoermer & Kreis 

Nitzschia homburgiensis Lange-Bertalot Stoermer et al. 

Nitzschia hungarica Grunow Stoermer & Kreis 

Nitzschia hustedtiana Salah Rushforth et al. 

Nitzschia hybrida Grunow Czamecki & Blinn 

Nitzschia ignorata Krasske Stoermer & Kreis 

Nitzschia impressa Hustedt Stoermer et al. 

Nitzschia incomptus Hohn & Hellerman Hohn & Hellerman 

Nitzschia inconspicua Grunow Stoermer et al. 

Nitzschia incurvata var. lorenziana R. Ross Stoermer et al. 

Nitzschia innominata Sovereign Stoermer et al. 

Nitzschia insecta Hustedt Stoermer & Kreis 

Nitzschia intermedia Hantzsch Stoermer & Kreis 

Nitzschia intermissa Hustedt Collins & Kalinsky 

Nitzschia interrupta (Reichelt) Hustedt Stoermer & Kreis 

Nitzschia invisitata Hustedt Dodd 

Nitzschia kittonii H. L. Smith Hohn & Hellerman 

Nitzschia kuetzingiana Hilse Stoermer & Kreis 

Nitzschia kuetzingiana var. exilis Grunow Stoermer & Kreis 

Nitzschia kuetzingioides Hustedt Stoermer et al. 

Nitzschia kurzii Rabenhorst Collins & Kalinsky 

Nitzschia laevis Hustedt Collins & Kalinsky 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 459 



Name Publication 



Nitzschia lacunanim Hustedt Stoermer & Kreis 

Nitzschia lacuum Lange-Bertalot Stoermer et al. 

Nitzschia lancettula O. Miiller Sovereign 

Nitzschia latens Hustedt Collins & Kalinsky 

Nitzschia lauenburgiana Hustedt Stoermer & Kreis 

Nitzschia lesbia Cholnoky Collins & Kalinsky 

Nitzschia levidensis (W. Smith) Grunow Cambum 

Nitzschia levidensis var. victoriae (Grunow) Cholnoky Collins & Kalinsky 

Nitzschia liebetruthii Rabenhorst Collins & Kalinsky 

Nitzschia linearis (Agardh) W. Smith Stoermer & Kreis 

Nitzschia linearis f. minuta Cleve & Moller 

Nitzschia linearis var. tenuis (Kiitzing) Grunow Stoermer & Kreis 

Nitzschia littoralis Grunow Patrick 

Nitzschia littoralis var. tergestina Grunow Czamecki & Blinn 

Nitzschia longissima (Brebisson) Ralfs Stoermer & Kreis 

Nitzschia longissima var. closterium (W. Smith) Van Heurck Stoermer & Kreis 

Nitzschia longissima var. reversa Grunow Stoermer & Kreis 

Nitzschia longissima f. parva Grunow Stoermer & Kxeis 

Nitzschia lorenziana Grunow Stoermer & Kreis 

Nitzschia lorenziana var. subtilis Grunow Cambum 

Nitzschia luzonensis Hustedt Stoermer & Kreis 

Nitzschia macilenta Gregory Stoermer & Kreis 

Nitzschia magnacarina Hohn & Hellerman Collins & Kalinsky 

Nitzschia manca Hustedt Hohn & Hellerman 

Nitzschia mediastalsis Hohn & Hellerman Hohn & Hellerman 

Nitzschia mediocris Hustedt Stoermer & Kreis 

Nitzschia microcephala Grunow Stoermer & Kreis 

Nitzschia microcephala var. elegantula Grunow Hohn & Hellerman 

Nitzschia migrans Cleve Collins & Kalinsky 

Nitzschia minuta Bleisch Camburn 

Nitzschia minutula Grunow Stoermer et al. 

Nitzschia mollis Hustedt Collins & Kalinsky 

Nitzschia monoensis Kociolek & Herbst Kociolek & Herbst 

Nitzschia montanestris Cambum Camburn 



Nitzschia nana Grunow Potapova & Charles 2003 

Nitzschia nereidis Cholnoky Collins & Kalinsky 1977 

Nitzschia obligata Archibald Collins & Kalinsky 1977 

Nitzschia obsidialis Hustedt Stoermer & Kreis 1978 

Nitzschia obtusa W. Smith Stoermer & Kreis 1978 

Nitzschia obtusa var. brevissima Grunow Dodd 1981 

Nitzschia obtusa var. nana Gmnow Camburn 1982 

Nitzschia obtusa var. scalpelliformis Gmnow Stoermer & Kreis 1978 

Nitzschia oregana Sovereign Clark & Rushforth 1977 

Nitzschia ovalis Amott Collins & Kalinsky 1977 

Nitzschia palea fKutzing) W. Smith Stoermer & Kreis 1978 

Nitzschia palea var. debilis (Kiitzing) Gmnow Patrick 1945 

Nitzschia palea var. sumatrana Hustedt Stoermer & Kreis 1978 

Nitzschia palea var. tenuirostris Gmnow Stoermer & Kreis 1978 

Nitzschia palea var. tropica Hustedt Collins & Kalinsky 1977 

Nitzschia paleacea Gmnow Stoermer & Kreis 1978 

Nitzschia paleaformis Hustedt Dmm 1981 

Nitzschia paleoides Hustedt Stoermer et al. 1999 

Nitzschia paradoxa (Gmelin) Gmnow Stoermer & Kreis 1978 

Nitzschia parvula W. Smith Stoermer & Kreis 1978 

Nitzschia parvula var. terricola Lund Collins & Kalinsky 1977 

Nitzschia paxillifer (O. F. Miiller) Heiberg Boyer 1927b 

Nitzschia perminuta Gmnow Camburn 1982 



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Nitzschia perspicillata Cambum Cambum 1982 

Nitzschia perspicua Sovereign Sovereign 1963 

Nitzschia pertica Hohn & Hellerman Hohn & Hellerman 1963 

Nitzschia perversa Grunow Dodd 1987 

Nitzschia philippinarum Hustedt Stoermer et al. 1999 

Nitzschia pilum Hustedt Hansmann 1973 

Nitzschia plana W. Smith Hansmann 1973 

Nitzschia plana var. americana Hustedt 1924 Hustedt 1924 

Nitzschia planctonica Hustedt Stoermer et al. 1999 

Nitzschia praetexta Nitzsch Whitford & Schumacher 1973 

Nitzschia pseudoamphioxys Hustedt Stoermer & Kreis 1978 

Nitzschia pseudobacata Cholnoky Collins & Kalinsky 1977 

Nitzschia pseudofonticola Hustedt Collins & Kalinsky 1977 

Nitzschia pseudohybrida Patrick & Roberts 1979 

Nitzschia pseudosinuata Hamilton & Laird Hamilton et al. 2001 

Nitzschia pseudostagnorum Hustedt Kaczmarska & Rushforth 1983 

Nitzschia pubens Cholnoky Collins & Kalinsky 1977 

Nitzschia pumila Hustedt Cambum 1982 

Nitzschia punctata (Grunow) Grunow Rushforth & Merkley 1988 

Nitzschia punctata var. elongata Grunow Rushforth & Merkley 1988 

Nitzschia punctata var. peragalli Halden Patrick 1961 

Nitzschia pura Hustedt Stoermer et al. 1999 

Nitzschia pusilla (Kiitzing) Grunow emend Lange-Bertalot Cambum 1982 

Nitzschia radicula Hustedt Stoermer et al. 1999 

Nitzschia radiosa Kutzing Czamecki & Blinn 1978 

Nitzschia rautenbachiae Cholnoky Cambum 1982 

Nitzschia recta Hantzsch Stoermer & Kreis 1978 

Nitzschia recta var. romana Collins & Kalinsky 1977 

Nitzschia rectiformis Hustedt Hohn 1961 

Nitzschia reimerii Kociolek & Herbst Kociolek & Herbst 1992 

Nitzschia regula Hustedt Patrick 1961 

Nitzschia reversa W. Smith Cambum 1982 

Nitzschia romana Grunow Stoermer & Kreis 1978 

Nitzschia romanoides Manguin Kalinsky 1983 

Nitzschia rostellata Hustedt Cambum 1982 

Nitzschia rufitorrentis Cholnoky Dodd 1987 

Nitzschia scalaris (Ehrenberg) W. Smith Stoermer & Kreis 1978 

Nitzschia scalaris var. undulata Wolle Tempere & Peragallo 1911 

Nitzschia semidesum Hohn & Hellerman Hohn & Hellerman 1963 

Nitzschia sentiformis Hohn & Hellerman Hohn & Hellerman 1963 

Nitzschia serpenticula Cholnoky Collins & Kalinsky 1977 

Nitzschia serpentiraphe Lange-Bertalot Lange-Bertalot 1993 

Nitzschia sicula var. migrans (Cleve) Hasle Stoermer & Kreis 1978 

Nitzschia sigma (Kutzing) W. Smith Stoermer & Kreis 1978 

Nitzschia sigma var. diminuta Gmnow Stoermer & Kreis 1978 

Nitzschia sigma var. rigida (Kutzing) Gmnow Stoermer et al. 1999 

Nitzschia sigma var. rigidula Gmnow Hohn & Hellerman 1963 

Nitzschia sigma var. sigmatella (Gregon,) Gmnow Stoermer & Kreis 1978 

Nitzschia sigmatella Gregory Boyer 1927b 

Nitzschia sigmaformis Hustedt Cambum 1982 

Nitzschia sigmoidea (Nitzsch) W. Smith Stoermer & Kreis 1978 

Nitzschia silicula Hustedt Czamecki et al. 1981 

Nitzschia siHcuia var. commutata Reimer Reimer 1966 

Nitzschia silicula var. migrans Patrick & Roberts 1979 

Nitzschia siliqua Archibald Collins & Kalinsky 1977 

Nitzschia sinuata W. Smith Stoermer & Kreis 1978 

Nitzschia sinuata var. delongnei (Gmnow) Lange-Bertalot Stoermer et al. 1999 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 461 



Name Publication 

Nitzschia sinuata var. tabellaria (Grunow) Grunow Stoermer & Kreis 1978 

Nitzschia sociabilis Hustedt Cambum 1982 

Nitzschia socialis Gregory Rushforth & Merkley 1988 

Nitzschia solita Hustedt Potapova & Charles 2002 

Nitzschia speciosa Hustedt Lowe 1972-1973 

Nitzschia spectabihs (Ehrenberg) Ralfs Collins & Kahnsky 1977 

Nitzschia spectabihs var. americana Grunow Boyer 1927b 

Nitzschia sphaerophora A. Cleve Stoermer & Kreis 1978 

Nitzschia spiculoides Hustedt Stoermer & Kreis 1978 

Nitzschia spiculum Hustedt Colhns & Kalinsky 1977 

Nitzschia stagnicola Rabenhorst Kalinsky 1983 

Nitzschia stagnorum Rabenhorst Stoermer & Kreis 1978 

Nitzschia steynii Cholnoky Collins & Kalinsky 1977 

Nitzschia stricta Hustedt Collins & Kalinsky 1977 

Nitzschia subacicularis Hustedt Stoermer & Kreis 1978 

Nitzschia subamphioxoides Hustedt Stoermer & Kreis 1978 

Nitzschia subcapitellata Hustedt Stoermer et al. 1999 

Nitzschia subconfmis Cholnoky Reimer 1982 

Nitzschia subhnearis Hustedt Stoermer & Kreis 1978 

Nitzschia subrostrata Hustedt Stoermer & Kreis 1978 

Nitzschia subrostratoides Cholnoky Collins & Kalinsky 1977 

Nitzschia subrostroides Cholnoky Stoermer et al. 1999 

Nitzschia subtilis (Kutzing) Grunow Stoermer & Kreis 1978 

Nitzschia subtilis var. paleacea Grunow Stoermer & Kreis 1978 

Nitzschia subvitrea Hustedt Dodd 1987 

Nitzschia suchlandtii Hustedt Potapova & Charles 2002 

Nitzschia supralitorea Lange-Bertalot Potapova & Charles 2003 

Nitzschia tabellaria Grunow Stoermer & Kreis 1978 

Nitzschia tarda Hustedt Stoermer & Kreis 1978 

Nitzschia tenuis W. Smith Stoermer & Kreis 1978 

Nitzschia terricola Lund Camburn et al. 1978 

Nitzschia thermalis (Ehrenberg) Auerswald Stoermer & Kreis 1978 

Nitzschia thermalis var. dubia Colhns & Kalinsky 1977 

Nitzschia thermalis var. intermedia Grunow Stoermer & ICreis 1978 

Nitzschia thermalis var. minor Hilse Stoermer & JCreis 1978 

Nitzschia tonoensis Foged Hansmann 1973 

Nitzschia tropica Hustedt Stoermer & Kreis 1978 

Nitzschia tryblionella Hantzsch Stoermer & Kreis 1978 

Nitzschia tryblionella var. calida (Grunow) Van Heurck Hohn & Hellerman 1963 

Nitzschia tryblionella var. debilis (Amott) A. Mayer Stoermer & Kreis 1978 

Nitzschia tryblionella var. levidensis (W. Smith) Grunow Stoermer & Kreis 1978 

Nitzschia tryblionella var. maxima Grunow Camburn et al. 1978 

Nitzschia trybhonella var. salinarum Grunow Hohn & Hellerman 1963 

Nitzschia tryblionella var. victoriae Grunow Stoermer & Kreis 1978 

Nitzschia umbilicata Hustedt Collins & Kalinsky 1977 

Nitzschia umbonata (Ehrenberg) Lange-Bertalot Dodd 1987 

Nitzschia valdestriata Aleem & Hustedt Camburn 1982 

Nitzschia valga Cholnoky Collins & Kalinsky 1977 

Nitzschia vermicularis (Kutzing) Grunow Stoermer & Kreis 1978 

Nitzschia vexans Grunow Stoermer & Kreis 1978 

Nitzschia vitrea Norman Stoermer & Kreis 1978 

Nitzschia vitrea var. major Tempere & Peragallo 1909 

Nitzschia vitrea var. recta (Hantzsch) Van Heurck Tilden 1894-1909 (#94) 

Nitzschia vitrea var. scaphiformis Wislough & Poretzky Dodd 1987 

Nitzschia vivax W. Smith Boyer 1927b 

Nitzschia vonhauseniae Cholnoky Collins & Kalinsky 1977 

Nitzschia volcanica Sovereign Sovereign 1958 



462 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Name Publication 

Nitzschia vulga Cholnoky Stoermer et al. 1999 

Nitzschia wipplingeri Cholnoky Collins & Kalinsky 1977 

Nitzschia woltereckii Hustedt Stoermer & Kreis 1978 

Nitzschia zuzulandica Cholnoky Collins & Kalinsky 1977 

Nupela Carolina Potapova & Clason 2003 Potapova et al. 2003 

Nupela neglecta Ponader, Lowe & Potapova 2003 Potapova et al. 2003 

Nupela neotropica Lange-Bertalot in Lange-Bertalot & Moser Siver et al. 2005 

Nupela paludigena (Scherer) Lange-Bertalot Lange-Bertalot 1993 

Nupela vitiosa (Schimanski) Lange-Bertalot Siver et al. 2005 

Nupela wellneri (Lange-Bertalot) Lange-Bertalot Potapova et al. 2003 

Odontella polymorpha Ehrenberg 1856 

Odontidium anceps Ehrenberg Patrick 1945 

Odontidium elongatum (Agardh) Kuntze Elmore 1922 

Odontidium hiemale (Roth) Heiberg Stoermer & Kreis 1978 

Odontidium hiemale var. mesodon (Ehrenberg) Grunow Patrick 1945 

Odontidium mesodon Mutzing Aubert 1895 

Odontidium mutabile W. Smith Stoermer & Kreis 1978 

Odontidium tabellaria Lewis Aubert 1895 

Odontidium vulgare (Bory) Elmore Elmore 1922 

Oestrupia bicontracta (0strup) Lange-Bertalot & Krammer Stoermer et al. 1999 

Oestrupia zachariasi (Reichelt) Stoermer & Yang Stoermer & Kreis 1978 

Oestrupia zachariasi var. undulata (Schultz) Stoermer & Yang Stoermer & Kreis 1978 

Opephora americana M. Peragallo Patrick & Reimer 1966 

Opephora ansata Hohn & Hellerman Stoermer & Kreis 1978 

Opephora martyi Heribaud Stoermer & Kreis 1978 

Opephora martyi var. capitata (Heribaud) Hustedt Sovereign 1958 

Opephora olsenii Moller Morales 2001 

Opephora pacifica Petit Prescott & Dillard 1979 

Opephora pinnata Ehrenberg Stoermer & Kreis 1978 

Opephora schulzi (Brockmann) Simons Czamecki et al. 1981 

Opephora swartzii (Grunow) Petit Prescott & Dillard 1979 

??Orthoseira dendroteres (Ehrenberg) Crawford Gaiser & Johansen 2000 

??Orthosira dickiei Thwaites Stoermer & Kreis 1978 

??Orthosira orichalcea Collins & Kalinsky 1977 

??Orthoseira roseana (Rabenhorst) O'Meara Hamilton et al. 1992 

O.xyneis binalis (Ehrenberg) Round in Round et al Siver et al. 2005 

Peronia fibula (Brebisson ex Kiitzing) Ross Patrick & Reimer 1966 

Peronia heribaudi Brun ex M. Peragallo in Heribaud Hamilton et al. 1992 

Peronia intermedium (H.L. Smith) Patrick Patrick & Reimer 1966 

Pinnularia abaujensis (Pantocsek) Ross Stoermer & Kreis 1978 

Pinnularia abaujensis var. iacustris Cambum & Charles Cambum & Charles 2000 

Pinnularia abaujensis var. linearis (Hustedt) Patrick Stoermer & Kreis 1978 

Pinnularia abaujensis var. rostrata (Patrick) Patrick Cambum 1982 

Pinnularia abaujensis var. subundulata (A. Mayer) Patrick Stoermer & Kreis 1978 

Pinnularia absita Hohn & Hellerman Patrick & Reimer 1966 

Pinnularia acrosphaeria W. Smith Stoermer & Kreis 1978 

Pinnularia acrosphaeria f. genuina Cleve Cleve 1895 

Pinnularia acrosohaeria var. laevis (M. Peragallo & Heribaud) Cleve Patrick & Reimer 1966 

Pinnularia acrosphaeria var. mrgidula Grunow ex Cleve Cambum 1982 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 463 



Name Publication 

Pinnularia acuminata W. Smith Patrick & Reimer 1966 

Pinnularia acuminata var. bielawski (Heribaud & Peragallo) Patrick Collins & Kalinsky 1977 

Pinnularia acuminata var. instabilis (A. Schmidt) Patrick Patrick & Reimer 1966 

Pinnularia acuminata var. interrupta (Clave) Patrick Patrick & Reimer 1966 

Pinnularia acuta Ehrenberg 1856 

Pinnularia aequalis Ehrenberg 1 856 

Pinnularia aequilateralis Patrick & Freese Stoermer 1964 

Pinnularia aestuartii Cleve Krammer 2000 

Pinnularia aestuartii var. interrupta (Hustedt) Cleve-Euler Bateman & Rushforth 1984 

Pinnularia affinis Ehrenberg 1856 

Pinnularia alabamae Krammer Krammer 2000 

Pinnularia alpina W. Smith Krammer 2000 

Pinnularia alpina var. elongata (M. Peragallo & Heribaud) Mills Patrick & Reimer 1966 

Pinnularia amblys Hohn & Hellerman Hohn & Hellerman 1963 

Pinnularia amphigomphus Ehrenberg 1856 

Pinnularia amphioxys Ehrenberg Stoermer & Kreis 1978 

Pinnularia amphisbaena Ehrenberg 1856 

Pinnularia amphistylus Ehrenberg Rushforth & Merkley 1988 

Pinnularia anglica Krammer Gaiser & Johansen 2000 

Pinnularia appendiculata (Agardh) Cleve Boyer 1927b 

Pinnularia aquilonaris Hohn & Hellerman Patrick & Reimer 1966 

Pinnularia balfouriana Patrick & Roberts 1979 

Pinnularia biceps Gregory Stoermer & Kreis 1978 

Pinnularia biceps var. minor (Petersen) Cleve-Euler Rushforth & Merkley 1988 

Pinnularia biceps f. petersenii Ross Stoermer & Kreis 1978 

Pinnularia biceps var. pusilla Cambum & Charles Cambum & Charles 2000 

Pinnularia bigibba Gaiser & Johansen Gaiser & Johansen 2000 

Pinnularia bihastata (A. Mann) Patrick Hansmann 1973 

Pinnularia bogotensis (Grunow) Cleve Boyer 1927b 

Pinnularia bogotensis var. undulata (Peragallo) Boyer Patrick & Reimer 1966 

Pinnularia borealis Ehrenberg Stoermer & Kreis 1978 

Pinnularia borealis var. brevicostata Hustedt Collins & Kalinsky 1977 

Pinnularia borealis var. caraccana (Ehrenberg) Brun Patrick & Reimer 1966 

Pinnularia borealis var. congolensis Zanon Loescher 1981 

Pinnularia borealis var. rectangularis Carlson Stoermer & Kreis 1978 

Pinnularia borealis var. scalaris (Ehrenberg) Rabenhorst Gaiser & Johansen 2000 

Pinnularia borealis var. subacuta Ehrenberg Patrick & Reimer 1966 

Pinnularia borealis var. truncata Ehrenberg Patrick & Reimer 1966 

Pinnularia boyeri Patrick Patrick & Reimer 1966 

Pinnularia bramanorum Ehrenberg 1 856 

Pinnularia brandelii Cleve Stoermer & Kreis 1978 

Pinnularia brauniana (Grunow ex Schmidt) F.W. Mills Hamilton et al. 1992 

Pinnularia braunii (Grunow) Cleve Stoermer & Kreis 1978 

Pinnularia braunii var. amphicephala (A. Mayer) Hustedt Stoermer & Kreis 1978 

Pinnularia braunii var. amphicephala f. subconica Venkataraman Cambum & Charles 2000 

Pinnularia brebissonii (Kutzingj Rabenhorst Stoermer & Kreis 1978 

Pinnularia brebissonii f. biundulata O. Miiller Lawson & Rushforth 1975 

Pinnularia brebissonii var. diminuta (Grunow) Cleve Stoermer & Kreis 1978 

Pinnularia brebissoni var. notata Heribaud & Peragallo Cleve 1895 

Pinnularia brevicostata Cleve Stoermer & Kreis 1978 

Pinnularia brevicostata var. leptostauron Cleve Tiffany & Britton 1952 

Pinnularia burkii Patrick Stoermer & Kreis 1978 

Pinnularia capitata Ehrenberg 1856 

Pinnularia cardinaliculis Cleve Stoermer & Kreis 1978 

Pinnularia cardinalis (Ehrenberg) W. Smith Stoermer & Kreis 1978 

Pinnularia carolinensis Ehrenberg 1 856 

Pinnularia castor Hohn & Hellerman Patrick & Reimer 1966 



464 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

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Name Publication 

Pinnularia caudata (Boyer) Patrick Cambum 1982 

Pinnularia cherryfieldiana Krammer Krammer 2000 

Pinnularia clevei Patrick Patrick 1945 

Pinnularia cocconeis Ehrenberg Stoermer & Kreis 1978 

Pinnularia conspicua (A. Schmidt) Cleve Patrick & Reimer 1966 

Pinnularia convexa Sovereign Patrick & Reimer 1966 

Pinnularia cooperi J.W. Bailey Ehrenberg 1856 

Pinnularia crucifera var. subrostrata A. Cleve Clark & Rushforth 1977 

Pinnularia cumvibia Hohn & Hellerman Hohn & Hellerman 1963 

Pinnularia cuneicephala (Mann) Patrick Patrick & Reimer 1966 

Pinnularia dactylus Ehrenberg Stoermer & Kreis 1978 

Pinnularia dactylus var. dariana (A. Schmidt) Cleve Boyer 1927b 

Pinnularia dactylus var. demerarae Cleve Boyer 1927b 

Pinnularia decurrans Ehrenberg Patrick & Reimer 1966 

Pinnularia dicephala (Ehrenberg) W. Smith Ehrenberg 1856 

Pinnularia digitus Ehrenberg 1856 

Pinnularia disphenia Ehrenberg 1856 

Pinnularia distinguenda Cleve Stoermer & Rreis 1978 

Pinnularia divergens W. Smith Stoermer & Kreis 1978 

Pinnularia divergens var. bacillaris (M. Peragallo) Mills Stoermer & Kreis 1978 

Pinnularia divergens var. decrescens (Grunow) Krammer Gaiser & Johansen 2000 

Pinnularia divergens var. elliptica (Grunow) Cleve Stoermer & Kreis 1978 

Pinnularia divergens var. parallela (Brun) Patrick Cambum 1982 

Pinnularia divergens var. schweinfurthii (A. Schmidt) Cleve Boyer 1927b 

Pinnularia divergens var. sublinearis Cleve Boyer 1927b 

Pinnularia divergens var. undulata (M. Peragallo & Heribaud) Hustedt Patrick & Reimer 1966 

Pinnularia divergentissima (Grunow) Cleve Patrick 1946 

Pinnularia divergentissima f. elongata Cleve-Euler Patrick 1946 

Pinnularia divergitissima var. subrostrata A. Cleve Siver et al. 2005 

Pinnularia doloma Hohn & Hellerman Patrick & Reimer 1966 

Pinnularia dubitabilis Hustedt Czamecki & Blinn 1978 

Pinnularia elliptica Ehrenberg 1856 

Pinnularia elongata Hustedt Hustedt 1934 

Pinnularia episcopalis Cleve Boyer 1927b 

Pinnularia episcopalis var. subelliptica A. Cleve Lawson & Rushforth 1975 

Pinnularia erratica var. fossilis Krammer Krammer 2000 

Pinnularia esox Cleve Boyer 1927b 

Pinnularia esoxiformes Fusey Gaiser & Johansen 2000 

Pinnularia falaiseana Krammer Gaiser & Johansen 2000 

Pinnularia fasciata Lagerstedt Stoermer & Kreis 1978 

Pinnularia ferroindulgentissima Czamecki & Cawley Czamecki & Cawley 1997 

Pinnularia flaminula A. Schmidt Patrick & Reimer 1966 

Pinnularia flexuosa Cleve Boyer 1927b 

Pinnularia flexuosa var. cuneata (Tempere & Peragallo) Mills Patrick & Reimer 1966 

Pinnularia flexuosa var. gibbosa Hustedt Hustedt 1934 

Pinnularia fluminea Patrick & Freese Lowe 1972-1973 

Pinnularia formica (Ehrenberg) Patrick Cambum 1982 

Pinnularia fossilis Krammer Krammer 2000 

Pinnularia gastmm Ehrenberg 1856 

Pinnularia gentilis (Donkin) Cleve Stoermer & Kreis 1978 

Pinnularia gibba Ehrenberg Stoermer & Kreis 1978 

Pinnularia gibba f. constricta Skvortzow Patrick & Reimer 1966 

Pinnularia gibba f. curta Rabenhorst Hohn & Hellerman 1963 

Pinnularia gibba var. gibba Hustedt Hustedt 1934 

Pinnularia gibba var. linearis Hustedt Stoermer & Kreis 1978 

Pinnularia gibba var. mesogongyla (Ehrenberg) Hustedt Reimer 1961 

Pinnularia gibba var. parva (Ehrenberg) Gmnow Stoermer & Kreis 1978 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 465 



Name Publication 

Pinnularia gibba var. rostrata Patrick Patrick 1945 

Pinnularia gibba f. subundulata A. Mayer Stoermer & Kreis 1978 

Pinnularia gibbiformis Krammer Krammer 2000 

Pinnularia gibbiformis var. floralensis Dute & Sullivan Dute et al. 2000 

Pinnularia gigas Ehrenberg Krammer 2000 

Pinnularia globiceps Gregory Stoermer & Kreis 1978 

Pinnularia globiceps var. krockii (Grunow) Cleve Stoermer & Kreis 1978 

Pinnularia gracillima Gregory Patrick & Reimer 1966 

Pinnularia gracilis Hustedt Lowe 1972-1973 

Pinnularia hannii Patrick Patrick 1946 

Pinnularia hemiptera Rabenhorst Stoermer & Kreis 1978 

Pinnularia hemiptera var. bielawski Heribaud & Peragallo Clark & Rushforth 1977 

Pirmularia hilseana Janisch Cambum 1982 

Pinnularia inaequalis Ehrenberg 1856 

Pinnularia instita Hohn & Hellerman Hohn & Hellerman 1963 

Pinnularia Integra Grunow in Cleve Cleve 1895 

Pinnularia intermedia (Lagerstedt) Cleve Stoermer & Kreis 1978 

Pinnularia intemipta W. Smith Stoermer & Kreis 1978 

Pinnularia intemipta f. bicapitata (Lagerstedt) Fritsch Patrick & Reimer 1966 

Pinnularia intemipta var. crassior (Grunow) Cleve Stoermer & Kreis 1978 

Pinnularia intemipta var. sinica Skvortzow Patrick & Reimer 1966 

Pinnularia iridis Ehrenberg 1 856 

Pinnularia isostauron (Ehrenberg) Cleve Patrick & Reimer 1966 

Pinnularia isselana Krammer Siver et al. 2005 

Pinnularia kasswingensis Ehrenberg 1856 

Pinnularia karelica Cleve Drum 1981 

Pinnularia kheuckeri Hustedt Collins & Kalinsky 1977 

Pinnularia kriegeriana Krasske emend Foged Stoermer et al. 1999 

Pinnularia krockii (Grunow) Cleve Stoermer et al. 1999 

Pinnularia kwacksii Cambum & Charles Cambum & Charles 2000 

Pinnularia lata (Brebisson) W. Smith Boyer 1927b 

Pinnularia lata var. amplissima Manguin Cambum 1982 

Pinnularia lata var. pachyptera (Ehrenberg) Meister Patrick & Reimer 1966 

Pinnularia lata var. rabenhorstii (Grunow) Cleve Boyer 1927b 

Pinnularia latevittata Cleve Patrick & Reimer 1966 

Pirmularia latevittata var. domingensis Cleve Stoermer & Kreis 1978 

Pinnularia latifascia Patrick Patrick 1946 

Pinnularia legumen (Ehrenberg) Ehrenberg Stoermer & Kreis 1978 

Pinnularia leptogongyla Ehrenberg Patrick & Reimer 1966 

Pinnularia leptosoma (Grunow) Cleve Stoermer & Kreis 1978 

Pinnularia leptosoma f. erlangensis A. Mayer Stoermer & Kreis 1978 

Pinnularia leptostigma Ehrenberg 1856 

Pinnularia longa Gregory Patrick & Reimer 1966 

Pinnularia lundii Hustedt Hohn & Hellerman 1963 

Pinnularia macilenta Ehrenberg Patrick 1946 

Pinnularia major (Kutzing) Rabenhorst Stoermer & Kreis 1978 

Pinnularia major var. asymmetrica Cleve Boyer 1927b 

Pinnularia major var. capitata Hustedt Hustedt 1934 

Pinnularia major var. heroina (A. Schmidt) Cleve Patrick & Reimer 1966 

Pinnularia major var. hustedti Meister Hansmann 1973 

Pinnularia major var. linearis Cleve Patrick & Reimer 1966 

Pinnularia major var. pulchella Boyer Boyer 1927b 

Pinnularia major var. subacuta (Ehrenberg) Cleve Patrick & Reimer 1966 

Pinnularia maior var. transversa (A. Schmidt) Cleve Cambum 1982 

Pinnularia major var. turgidula Cleve Patrick & Reimer 1966 

Pinnularia makahana Sovereign Patrick & Reimer 1966 

Pinnularia martyi Lauby Prescott & Dillard 1979 



466 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Name Publication 

Pinnularia megaloptera Ehrenberg 1856 

Pinnularia mesogonglya Ehrenberg Cambum 1982 

Pinnularia mesolepta (Ehrenberg) W. Smith Stoermer & Kreis 1978 

Pinnularia mesolepta var. angusta Cleve Patrick & Reimer 1966 

Pinnularia mesolepta var. stauroneiformis (Grunow) Cleve Patrick & Reimer 1966 

Pinnularia mesolepta var. turbulenta Cleve-Euler Clark & Rushforth 1977 

Pinnularia mesotyla Ehrenberg Patrick & Reimer 1966 

Pinnularia microstauron (Ehrenberg) Cleve Stoermer & Kreis 1978 

Pinnularia microstauron var. adirondackensis Cambum & Charles Cambum & Charles 2000 

Pinnularia microstauron var. biundulata O. Miiller Stoermer & Kreis 1978 

Pinnularia microstauron var. lunicus Cambum & Charles Cambum & Charles 2000 

Pinnularia microstauron var. nonfasciata Krammer Krammer 2000 

Pinnularia molaris (Gmnow) Cleve Boyer 1927b 

Pinnularia molaris var. asiatica Skvortzow Cambum et al. 1978 

Pinnularia montgomeryana Krammer Krammer 2000 

Pinnularia moralis (Gmnow) Cleve Stoermer & Kreis 1978 

Pinnularia mormonomm (Gmnow) Boyer Boyer 1927b 

Pinnularia neglecta (A. Mayer) A. Berg Reimer 1990 

Pinnularia neomajor Krammer Siver et al. 2005 

Pinnularia nobilis (Ehrenberg) W. Smith Stoermer & Kreis 1978 

Pinnularia nodosa (Ehrenberg) W. Smith Stoermer & Kreis 1978 

Pinnularia nodosa var. constricta f. tmncata Fusey Cambum & Charles 2000 

Pinnularia nodosa var. pseudogracillima (May) A. Cleve Clark & Rushforth 1977 

Pinnularia nodosa var. robusta (Foged) Krammer Siver et al. 2005 

Pinnularia notabilis Krammer Gaiser & Johansen 2000 

Pinnularia nubila Sovereign Patrick & Reimer 1966 

Pinnularia obscura Krasske Stoermer & Kreis 1978 

Pinnularia obtusa Ehrenberg Ehrenberg 1856 

Pinnularia obtusa Sovereign Patrick & Reimer 1966 

Pinnularia ohiensis Ehrenberg Ehrenberg 1856 

Pinnularia oregonica Ehrenberg 1856 

Pinnularia oxylepta Ehrenberg Ehrenberg 1856 

Pinnularia oxy trachea Ehrenberg Ehrenberg 1856 

Pinnularia palousiana Sovereign Patrick & Reimer 1966 

Pinnularia parallela Bmn Boyer 1 927b 

Pinnularia parva Gregory Boyer 1927b 

Pinnularia parvula (Rails) Cleve-Euler Patrick & Reimer 1966 

Pinnularia paulensis Gmnow in Cleve Boyer 1927b 

Pinnularia peregrina Ehrenberg 1856 

Pinnularia permagna J.W. Bailey Ehrenberg 1856 

Pinnularia placentula Ehrenberg 1856 

Pinnularia platycephala f. omata Sovereign Patrick & Reimer 1966 

Pinnularia platysoma Ehrenberg 1856 

Pinnularia pluviana Sovereign Krammer 2000 

Pinnularia podzorski Krammer Krammer 2000 

Pinnularia pogoii Scherer Scherer 1988 

Pinnularia polyonca (Brebisson) W. Smith Boyer 1927b 

Pinnularia pulchella (Boyer) Kranmier Krammer 2000 

Pinnularia pulchra 0stmp Clark & Rushforth 1977 

Pinnularia radiosa W. Smith Stoermer & Kreis 1978 

Pinnularia rivularis Hustedt Patrick & Reimer 1966 

Pinnularia mpestris Hantzsch Stoermer & Kreis 1978 

Pinnularia mttneri Hustedt Stoermer & Kreis 1978 

Pinnularia sabae Ehrenberg Patrick & Reimer 1966 

Pinnularia schweinfurthii (A. Schmidt) Patrick Patrick & Reimer 1966 

Pinnularia secemenda (A. Schmidt) Cleve Boyer 1927b 

Pinnularia semen Ehrenberg 1 856 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 467 



Name Publication 

Pinnularia semicniciata A. Cleve Stoermer & Kreis 1978 

Pinnularia signata Ehrenberg 1 856 

Pinnularia silicula Ehrenberg 1856 

Pinnularia sillimanorum Ehrenberg Boyer 1927b 

Pinnularia singularis (A. Schmidt) Cleve Patrick & Reimer 1966 

Pinnularia sirokiana Ehrenberg 1 856 

Pinnularia sociahs (T.C. Palmer) Hustedt Stoermer & Kreis 1978 

Pinnularia stauroneis Ehrenberg Rushforth & Merkley 1988 

Pinnularia stauroptera (Grunow) Cleve Boyer 1927b 

Pinnularia stauroptera var. interrupta Cleve Patrick & Reimer 1966 

Pinnularia stauroptera var. semicniciata Cleve Patrick & Reimer 1966 

Pinnularia stomatophora Grunow Boyer 1927b 

Pinnularia stromatophora (Grunow) Cleve Stoermer & Kreis 1978 

Pinnularia streptoraphe Cleve Boyer 1927b 

Pinnularia streptoraphe var. muscicola Skvortzow Reimer 1990 

Pinnularia striata Ehrenberg 1 856 

Pinnularia stricta Hustedt Hohn & Hellerman 1963 

Pinnularia subanglica Krammer Siver et al. 2005 

Pinnularia subcapitata Gregory Stoermer & Kreis 1978 

Pinnularia subcapitata var. hilseana (Janisch) O. Miiller Collins & Kalinsky 1977 

Pinnularia subcapitata var. hybrida (Grunow) Frenguelli Hohn & Hellerman 1963 

Pinnularia subcapitata var. lapponica A. Cleve Clark & Rushforth 1977 

Pinnularia subcapitata var. paucistriata (Grunow) Cleve Stoermer & Kreis 1978 

Pinnularia subcapitata var. stauroneiformis Van Heurck Patrick 1945 

Pinnularia subgibba var. gracilis Gaiser & Johansen Gaiser & Johansen 2000 

Pinnularia subgibba var. hustedtii Krammer Gaiser & Johansen 2000 

Pinnularia subgibba var. lanceolata Gaiser & Johansen Gaiser & Johansen 2000 

Pinnularia subgibba var. sublinearis Krammer Krammer 2000 

Pinnularia sublinearis (Grunow) Cleve Patrick 1945 

Pinnularia subnodosa Hustedt Hustedt 1934 

Pinnularia subpalousiana Sovereign Patrick & Reimer 1966 

Pinnularia subrostrata A. Cleve Stoermer & Kreis 1978 

Pinnularia subsolaris (Grunow) Cleve Sovereign 1958 

Pinnularia substomatophora Hustedt Stoermer & Kreis 1978 

Pinnularia sudetica Hilse Stoermer & Kreis 1978 

Pinnularia sudetica var. brittanica (Grunow) Krammer Krammer 2000 

Pinnularia sudetica var. commutata (Grunow) Cleve-Euler Johansen et al. 1983 

Pinnularia suecia Ehrenberg 1 856 

Pinnularia superba Cleve-Euler Clark & Rushforth 1977 

Pinnularia tabellaria Ehrenberg Stoermer & Kreis 1978 

Pinnularia tabellaria var. stauroneiformis Van Heurck Boyer 1927b 

Pinnularia tenuis Gregory Stoermer & Kreis 1978 

Pinnularia tenuis var. interrupta (Font.) A. Cleve Stoermer & Kreis 1978 

Pinnularia termes (Ehrenberg) A. Schmidt Boyer 1927b 

Pinnularia termitina (Ehrenberg) Patrick Stoermer & Kreis 1978 

Pinnularia tibetana Hustedt Stoermer & Kreis 1978 

Pinnularia titusiana Hagelstein Lowe 1972-1973 

Pinnularia torta (A. Mann) Patrick Stoermer & Kreis 1978 

Pinnularia transversa (W. Smith) A. Mayer Gaiser & Johansen 2000 

Pinnularia trigonocephala Cleve Stoermer & Kreis 1978 

Pinnularia turfosiphila Gaiser & Johansen Gaiser & Johansen 2000 

Pinnularia tumerae Cambum & Charles Camburn & Charies 2000 

Pinnularia umbrosa Sovereign Patrick & Reimer 1966 

Pinnularia undula (Schumann) Krammer Krammer 2000 

Pinnularia undula var. major (A. Schmidt) Krammer Krammer 2000 

Pinnularia undula var. mesoleptiformis Krammer Krammer 2000 

Pinnularia undulata Gregory Stoermer & Kreis 1978 



468 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

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Name Publication 

Pinnularia undulata var. subundulata Grunow Stoermer & Kreis 1978 

Pinnularia ventricosa Hustedt Hustedt 1934 

Pinnularia vespa Ehrenberg Patrick & Reimer 1966 

Pinnularia viridiformes Krammer Gaiser & Johansen 2000 

Pinnularia viridis (Nitzsch) Ehrenberg Stoermer & Kreis 1978 

Pinnularia viridis var. commutata (Grunow) Cleve Stoermer & Kreis 1978 

Pinnularia viridis var. caudata Boyer Boyer 1927b 

Pinnularia viridis var. commutata (Grunow) Cleve Boyer 1927b 

Pinnularia viridis var. elliptica Meister Patrick 1945 

Pinnularia viridis var. fallax Cleve Boyer 1927b 

Pinnularia viridis var. minor Cleve Collins & Kalinsky 1977 

Pinnularia viridis var. rupestris (Hantzsch) Cleve Hohn 1951 

Pinnularia viridis var. sudetica (Hilse) Hustedt Patrick 1945 

Pinnularia viridula Ehrenberg 1856 

Pinnularia wisconsinensis Cambum & Charles Cambum & Charles 2000 

Pinnularia woerthensis (A. Mayer) Krammer Gaiser & Johansen 2000 

Placoneis abiskoensis (Hustedt) Lange-Bertalot & Metzeltin Johansen et al. 2004 

Placoneis anglica (Ralfs in Pritchard) Lowe in Johansen et al Johansen et al. 2004 

Placoneis clementoides (Hustedt) E.J. Cox Stoermer et al. 1999 

Placoneis dementis (Grunow) E.J. Cox Stoermer et al. 1999 

Placoneis dicephala (W. Smith) Mereschkowsky Stoermer et al. 1999 

Placoneis elginensis (Gregory) Stoermer et al. 1999 

Placoneis exigua (Gregory) Mereschkowsky Stoermer et al. 1999 

Placoneis gastrum (Ehrenberg) Mereschkowsky Stoermer et al. 1999 

Placoneis lata (M. Peragallo) Lowe in Johansen et al Johansen et al. 2004 

Placoneis neglecta (Krasske) Lowe in Johansen et al Johansen et al. 2004 

Placoneis placentula (Ehrenberg) Heinzerling Stoermer et al. 1999 

Placoneis placentula f. rostrata (Mayer) Bukhtiyarova Stoermer et al. 1999 

Placoneis placentula var. rostrata (A. Mayer) Andresen et al Andresen et al. 2000 

Plagiotropis arizonica Czamecki & Blinn Czarnecki et al. 1981 

Plagiotropis lepidoptera (Gregory) Reimer Stoermer et al. 1999 

Plagiotropis lepidoptera var. proboscidea (Cleve) Reimer Cambum 1982 

Plagiotropis vitrea (W. Smith) Grunow Rushforth & Squires 1985 

Plagiotropis vitrea var. scaligera (Grunow ex Cleve & Grunow) Peragallo Rushforth & Squires 1985 

Planothidium apiculatum (Patrick) Lowe in Johansen et al Johansen et al. 2004 

Planothidium calcar (Cleve) Bukhtiyarova & Round Stoermer et al. 1999 

Planothidium delicatulum (Kutzing) Round & Bukhtiyarova Stoermer et al. 1999 

Planothidium dubium (Grunow) Round & Bukhtiyarova Stoermer et al. 1999 

Planothidium ellipticum (A. Cleve) Round & Bukhtiyarova Stoermer et al. 1999 

Planothidium fossile (Tempere & Peragallo) Lowe in Johansen et al Johansen et al. 2004 

Planothidium frequentissimum (Lange-Bertalot) Round & Bukhtiyarova Siver et al. 2005 

Planothidium hauckianum (Grunow) Round & Bukhtiyarova Stoermer et al. 1999 

Planothidium hauckianum var. rostratum (Schulz) Andresen et al Andresen et al. 2000 

Planothidium lanceolatum (Brebisson) Round & Bukhtiyarova Stoermer et al. 1999 

Planothidium lanceolatum var. bimaculatum (Hustedt) Andresen et al Andresen et al. 2000 

Planothidium lanceolatum var. genuinum (A. Mayer) Andresen et al Andresen et al. 2000 

Planothidium lanceolatum var. omissum (Reimer) Andresen et al Andresen et al. 2000 

Planothidium oestrupii (H. Bachmann & A. Cleve) Round & Bukhtiyarova Stoermer et al. 1999 

Planothidium peragallii (Brun & Heribaud) Round & Bukhtiyarova Stoermer et al. 1999 

Planothidium peragalli var. fossilum (Tempere & Peragallo) Andresen et al Andresen et al. 2000 

Planothidium peragalli var. parvulum (Patrick) Andresen et al Andresen et al. 2000 

Piayaensis circumfimbriata Spaulding & Kociolek Spaulding et al. 2002 

Playaensis furtiva Spaulding & Kociolek Spaulding et al. 2002 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 469 

Name Publication 

Pleurosira laevis (Ehrenherg) Compere Stoermer et al. 1999 

Pleurosigma acuminatum (Kiitzing) Grunow Stoermer & Kreis 1978 

Pleurosgima angulatum (Quekett) W. Smith Dodd 1987 

Pleurosigma attenuatum W. Smith Stoermer & Kreis 1978 

Pleurosigma australe Grunow Patrick & Reimer 1966 

Pleurosigma boyeri Keeley Keeley 1926 

Pleurosigma delicatulum W. Smith Stoermer & Kreis 1978 

Pleurosigma delicatulum var. americana Cleve Patrick & Reimer 1966 

Pleurosigma elongatum W. smith Patrick & Reimer 1966 

Pleurosigma eximium (Thwaites) Cleve & Grunow Stoermer & Kreis 1978 

Pleurosigma fasciolata W. Smith Ehrenberg 1856 

Pleurosigma formosum Sterrenburg 1991 

Pleurosigma kuetzingii Grunow Stoermer & Kreis 1978 

Pleurosigma normani Patrick & Reimer 1966 

Pleurosigma obscurum W. Smith Stoermer & Kreis 1978 

Pleurosigma obmsatum Sullivant & Wormley Collins & Kalinsky 1977 

Pleurosigma paradoxum Peragallo Patrick & Reimer 1966 

Pleurosigma salinarum Grunow Potapova & Charles 2002 

Pleurosigma salinarum var. boyeri (Keeley) Reimer Patrick & Reimer 1966 

Pleurosigma scalprum Grunow Cleve & Moller 1879 

Pleurosigma sciotense Sullivan Stoermer & Kreis 1978 

Pleurosigma spencerii Grunow Stoermer & Kreis 1978 

Pleurosigma spenceri (Quekett) W. Smith Tilden 1894-1909 (#90) 

Pleurosigma strigosum W. Smith Patrick & Reimer 1966 

Pleurosigma wansbeckii Donkin Cleve & Moller 1879 

Pleurosigma wormleyi Sullivan Stoermer & Kreis 1978 

Pleurostauron acuta Rabenhorst Tempere & Peragallo 1909 

Pleurostauron acuta var. robusta Tempere & Peragallo 1909 

Pleurostauron javanica Grunow Tempere & Peragallo 1911 

Pleurostauron smithii Grunow Tempere & Peragallo 1911 

Psammothidium abundans f. rosenstockii (Lange-Bertalot) Bukhtiyarova Stoermer et al. 1999 

Psammothidium altaicum (Poretzky) Bukhtiyarova & Round Stoermer et al. 1999 

Psammothidium bioreti (Germain) Bukhtiyarova & Round Stoermer et al. 1999 

Psammothidium daonense (Lange-Bertalot) Lange-Bertalot Siver et al. 2005 

Psammothidium didymum (Hustedt) Bukhtiyarova & Round Stoermer et al. 1999 

Psammothidium helveticum (Hustedt) Bukhtiyarova Potapova & Charles 2002 

Psanunothidium lauenburgianum (Hustedt) Round & Bukhtiyarova Potapova & Charles 2003 

Psammothidium levanderi (Hustedt) Bukhtiyarova & Round Stoermer et al. 1999 

Psammothidium marginulatum (Grunow) Bukhtiyarova & Round Stoermer et al. 1999 

Psammothidium sacculum (Carter) Bukhtiyarova in Bukhtiyarova & Round Siver et al. 2005 

Psammothidium subatomoides (Hustedt) Bukhtiyarova & Round Stoermer et al. 1999 

Psammothidium ventralis (Krasske) Bukhtiyarova & Round Stoermer et al. 1999 

Pseudoeunotia lunaris Ehrenberg Stoermer & Kreis 1978 

Pseudostaurosira brevistriata (Grunow in Van Heurck) Williams & Round Stoermer et al. 1999 

Pseudostaurosira brevistriata var. binodis (Pantocsek) Andresen et al Andresen et al. 2000 

Pseudostaurosira brevistriata var. capitata (Heribaud) Andresen et al Andresen et al. 2000 

Pseudostaurosira brevistriata var. inflata (Pantocsek) Edlund Stoermer et al. 1999 

Pseudostaurosira clavatum Morales Morales 2002 

Pseudostaurosira construens var. binodis Stoermer et al. 1999 

Pseudostaurosira neoelliptica (Witkowski) Morales Morales 2002 

Pseudostaurosira parasitica (W. Smith) Morales Morales 2003 

Pseudostaurosira parasitica var. subconstricta (Grunow) Morales Morales 2003 



470 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Name Publication 

Pseudostaurosira pseudoconstruens (Marciniak) Williams & Round Morales 2001 

Pseudostaurosira trainorii Morales Morales 2001 

Pseudostaurosiropsis connecticutensis Morales Morales 2001 

Pseudostaurosiropsis geocollegarum (Witkowski & Lange-Bertalot) Morales Morales 2002 

Reimeria sinuata (Gregory) Kociolek & Stoermer Stoermer et al. 1999 

Reimeria sinuata f. antiqua (Gregory) Kociolek & Stoermer Stoermer et al. 1999 

Rhizosolenia eriensis H.L. Smith Stoermer & Kreis 1978 

Rhizosolenia eriensis var. morsa West & West Gaufm et al. 1976 

Rhizosolenia eriensis var. pusilla Wolosz. In Schroeder Stoermer et al. 1999 

Rhizosolenia eriensis var. zachariasi (Brun) Playfair Stoermer et al. 1999 

Rhizosolenia gracilis H.L. Smith Stoermer & Kreis 1978 

Rhizosolenia longiseta Zacharias Stoermer & Kreis 1978 

Rhizosolenia minima Levan Czamecki et al. 1981 

Rhizosolenia rothii (Ehrenberg) Grunow Prescott & Dillard 1979 

Rhizosolenia stagnalis Zacharias Prescott & Dillard 1979 

Rhoicosphenia curvata (Kutzing) Grunow Stoermer & Kreis 1978 

Rhoicosphenia curvata var. gracilis Tempere & Peragallo 1912 

Rhoicosphenia curvata var. major Cleve Sovereign 1958 

Rhoicosphenia curvata var. minor Patrick 1968 

Rhoicosphenia curvata var. subacuta M. Schmidt Stoermer & Kreis 1978 

Rhopalodia argus W. Smith Stoermer & Kreis 1978 

Rhopalodia brebissonii Krammer Gaiser & Johansen 2000 

Rhopalodia gibba (Ehrenberg) O. Muller Stoermer & Kreis 1978 

Rhopalodia gibba var. parallela (Ehrenberg) O. Muller Reimer 1961 

Rhopalodia gibba var. ventricosa (Kiitzing) Peragallo & Peragallo Stoermer & Kreis 1978 

Rhopalodia gibberula (Ehrenberg) O. Muller Stoermer & Kreis 1978 

Rhopalodia gibberula var. producta (Grunow) O. Muller Patrick & Reimer 1975 

Rhopalodia gibberula var. protracta Grunow Patrick & Reimer 1975 

Rhopalodia gibberula var. rupestris (W. Smith) O. Muller Patrick & Reimer 1975 

Rhopalodia gibberula var. vanheurckii O. Muller Cambum 1982 

Rhopalodia musculus (Kutzing) O. Muller Cambum 1982 

Rhopalodia musculus var. constricta (Brebisson) Peragallo & Peragallo Rushforth & Merkley 1988 

Rhopalodia parallela (Grunow) O. Muller Stoermer & Kreis 1978 

Rhopalodia ventricosa (Kutzing) 0strup Prescott & Dillard 1979 

Rossithidium linearis (W. Smith) Round & Bukhtiyarova Stoermer et al. 1999 

Rossithidium pusillum (Grunow ) Round & Bukhtiyarova Stoermer et al. 1999 

Rouxia califomica var. minuta A. Cleve Stoermer & Kreis 1978 

Sarcophagodes delicatula Morales 2002 Morales 2002 

Sceptroneis fibula (Brebisson) Schutt Elmore 1922 

Sceptroneis pacifica (Grunow) Elmore Collins & Kalinsky 1977 

Schizonema viridulum Brebisson Stoermer & Kreis 1978 

Schizonema vulgare Thwaites Stoermer & Kreis 1978 

Scoliopleura campylogramma (Ehrenberg) Rabenhorst Stoermer et al. 1999 

Scoliopleura peisonis Grunow Patrick & Reimer 1966 

Scoliopleura tumida (Brebisson) Rabenhorst Elmore 1922 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 471 



Name Publication 

Sellaphora americana (Ehrenberg) D.G Mann Stoermer et al. 1999 

Sellaphora bacilarioides (Grunow) Andresen et al Andresen et al. 2000 

SeUaphora bacilliformis (Grunow) Mereschkowsky Stoermer et al. 1999 

Sellaphora bacillum (Ehrenberg) D.G. Mann Stoermer et al. 1999 

Sellaphora disjuncta (Hustedt) D.G Mann Hamilton et al. 1992 

Sellaphora laevissima (Kiitzing) D.G Mann Stoermer et al. 1999 

Sellaphora mutata (Krasske) Lange-Bertalot Stoermer et al. 1999 

Sellaphora nyassensis (O. Miiller) D.G Mann Stoermer et al. 1999 

Sellaphora nyassensis f. minor (O. Miiller) Andresen et al Andresen et al. 2000 

SeUaphora parapupula Lange-Bertalot Stoermer et al. 1999 

Sellaphora pupula (Kiitzing) Mereschkowsky Stoermer et al. 1999 

Sellaphora pupula var. elliptica (Hustedt) Poulin Stoermer et al. 1999 

Sellaphora pupula var. mutata (Krasske) Poulin Stoermer et al. 1999 

Sellaphora pupula f. rostrata (Hustedt) Bukhtiyarova Stoermer et al. 1999 

SeUaphora pupula var. rectangularis (Gregory) Mereschkowsky Stoermer et al. 1999 

SeUaphora rectangularis (Gregory) Lange-Bertalot & Metzeltin Stoermer et al. 1999 

Sellaphora rostrata (Hustedt) Johansen in Johansen et al Johansen et al. 2004 

SeUaphora seminulum (Grunow) D.G Mann Stoermer et al. 1999 

Sellaphora wummensis Johansen in Johansen et al Johansen et al. 2004 

Sellaphora vitabunda (Hustedt) D.G Mann Stoermer et al. 1999 

Semiorbis hemicyclus (Ehrenberg) Patrick Patrick & Reimer 1966 

Simonsenia delongei (Grunow) Lange-Bertalot Potapova & Charles 2002 

Skeletonema costatum (GreviUe) Cleve Stoermer & Kreis 1978 

Skeletonema potamos (Weber) Hasle Stoermer & Kreis 1978 

Skeletonema subsalsum (A. Cleve) Bethge Stoermer & Kreis 1978 

Sphinctocystis eUiptica (Kiitzing) Kuntze Stoermer & Kreis 1978 

Sphinctocystis librilis (Ehrenberg) Hassal Stoermer & Kreis 1978 

Stauroforma exiguiformis (Lange-Bertalot) Flower Morales 2001 

Stauroforma inermis Flower, Jones & Round Morales 2001 

Stauroneis acuta W. Smith Stoermer & Kreis 1978 

Stauroneis acuta var. major Tempere & Peragallo 1909 

Stauroneis acuta var. terryana Tempere ex Cleve Patrick & Reimer 1966 

Stauroneis acutiuscula M. Peragallo & Heribaud Stoermer & Kreis 1978 

Stauroneis agrestris Peterson Stoermer & Kreis 1978 

Stauroneis alabamae Heiden Stoermer & Kreis 1978 

Stauroneis alabamae var. angulata Heiden Hohn 195 1 

Stauroneis alabamae var. rostrata Heiden Heiden 1 903 

Stauroneis americana Heiden Boyer 1927b 

Stauroneis amphilepta Ehrenberg Patrick & Reimer 1966 

Stauroneis amphioxys Gregory Stoermer & Kreis 1978 

Stauroneis amphioxys var. obtusa Hendey Kalinsky 1983 

Stauroneis anceps Ehrenberg Stoermer & Kreis 1978 

Stauroneis anceps var. americana Reimer Stoermer & Kreis 1978 

Stauroneis anceps var. amphicephala (Kutzing) Cleve Patrick 1945 

Stauroneis anceps var. birostris (Ehrenberg) Cleve Patrick & Reimer 1966 

Stauroneis anceps var. capitata M. Peragallo Tempere & Peragallo 1908 

Stauroneis anceps var. elongata Tempere & Peragallo Tempere & Peragallo 1911 

Stauroneis anceps var. fossilis Cleve Tempere & Peragallo 1912 

Stauroneis anceps var. gracilis (Ehrenberg) Brun Patrick 1945 

Stauroneis anceps var. hyalina Brun & M. Peragallo Stoermer & Kreis 1978 

Stauroneis anceps var. linearis (Ehrenberg) Van Heurck Patrick 1945 



472 



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Name 

Staurone 
Staurone 
Staurone: 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Staurone 
Stuarone 
Staurone 



Publication 

s anceps var. nobilis Tempere & Peragallo 1909 

s anceps var. prominula Grunow Hohn 195 1 

s anceps var. siberica Grunow Stoermer & Kreis 1978 

s anceps var. subrostrata Gaiser & Johansen Gaiser & Johansen 2000 

s anceps f. gracilis Rabenhorst Stoermer & Kreis 1978 

s anceps f. linearis (Ehrenberg) Hustedt Stoermer & Kreis 1978 

s arctica A. Cleve Bateman & Rushforth 1984 

s baileyi Ehrenberg Boyer 1927b 

s baconiana Stodder Boyer 1927b 

s binodis Ehrenberg Patrick & Reimer 1966 

s birostris Ehrenberg Patrick & Reimer 1966 

s borrichii (Petersen) Lund Dodd 1987 

s borrichii f. subcapitata Peterson Collins & Kalinsky 1977 

s bovbjergii Reimer Reimer 1990 

s brunii M. Peragallo & Heribuad Tempere & Peragallo 1913 

s crucicula (Grunow) Boyer Boyer 1927b 

s dicephala Ehrenberg Patrick & Reimer 1966 

s dilatata Ehrenberg Stoermer & Kreis 1978 

s dilatata f. baicalensis Skvortzow & Meyer Stoermer & Kreis 1978 

s euglypta Ehrenberg Patrick & Reimer 1966 

s eurysoma Ehrenberg Patrick & Reimer 1966 

s fenestra Ehrenberg Patrick & Reimer 1966 

s fluminea Patrick & Freese Stoermer & Kreis 1978 

s frickei var. angusta Boyer Boyer 1927b 

s gallica M. Peragallo Patrick & Reimer 1966 

s gracilior (Rabenhorst) Reichardt Siver et al. 2005 

s gracihs Ehrenberg Stoermer & Kreis 1978 

s gregorii Ralfs Stoermer & Kreis 1978 

s ignorata Hustedt Collins & Kalinsky 1977 

s ignorata var. rupestris (Skvortzow) Reimer Hansmann 1973 

s javanica Grunow Cleve 1 894 

s kriegeri Patrick Stoermer & Kreis 1978 

s kriegeri f. undulata Hustedt Stoermer & Kreis 1978 

s lanceolata Kutzing Tempere & Peragallo 1911 

s lauenburgiana Hustedt Stoermer & Kreis 1978 

s legleri Hustedt Kaczmarska & Rushforth 1983 

s legumen (Ehrenberg) Kiitzing Boyer 1927b 

s linearis Ehrenberg Stoermer & Kreis 1978 

s lineolata Ehrenberg Patrick & Reimer 1966 

s livingstonii Reimer Stoermer & Kreis 1978 

s lundii Hustedt Clark & Rushforth 1977 

s macrocephala Kiitzing Patrick & Reimer 1966 

s maculata Ehrenberg 1856 

s montana Krasske Collins & Kalinsky 1977 

s muriella f. linearis Lund Rushforth & Squires 1985 

s nana Hustedt Patrick & Reimer 1966 

s neohyalina Lange-Bertalot Stoermer et al. 1999 

s nobilis Schumann Patrick & Reimer 1966 

s nobilis var. alabamae (Heiden) Cleve-Euler Cambum 1982 

s nobilis var. baconiana (Stodder) Reimer Stoermer & Kreis 1978 

s nobilis var. gracilis Kobayasi Cambum & Charles 2000 

s norvegica Hustedt Patrick & Reimer 1966 

s obtusa Lagerstedt Stoermer & Kreis 1978 

s obtusa var. catarinensis Krasske Dodd 1987 

s palustris Hustedt Drum 1981 

s parsula (Grunow) Cleve Boyer 1927b 

s phoenicenteron (Nitzsch) Ehrenberg Stoermer & Kreis 1978 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 473 



Name Publication 

Stauroneis phoenicenteron var. amphilepta (Ehrenberg) Cleve Tiffany & Britton 1952 

Stauroneis phoenicenteron var. baileyi Ehrenberg Cleve 1894 

Stauroneis phoenicenteron var. brevis Dippel Stoermer & Kreis 1978 

Stauroneis phoerucenteron var. brunii (Peragallo & Heribaud) Voigt Collins & Kalinsky 1977 

Stauroneis phoenicenteron var. genuina Cleve Cleve 1894 

Stauroneis phoenicenteron f. gracilis (Ehrenberg) Hustedt Stoermer & Kreis 1978 

Stauroneis phoenicenteron var. intermedia (Dippel) A. Cleve Stoermer et al. 1999 

Stauroneis phoenicenteron f. lanceolata (Kutzing) Brun Stoermer & Kreis 1978 

Stauroneis phoenicenteron f. linearis (Ehrenberg) Hustedt Reimer 1990 

Staiu^oneis phyllodes Ehrenberg Stoermer & Kreis 1978 

Stauroneis platalea Ehrenberg Patrick & Reimer 1966 

Stauroneis platysoma Ehrenberg Kalinsky 1983 

Stauroneis producta Grunow Stoermer & Kreis 1978 

Stauroneis pteroidea Ehrenberg Tempere & Peragallo 1908 

Stauroneis pulchella W. Smith Tilden 1894-1909 (#367) 

Stauroneis pygmaea Krieger Dodd 1981 

Stauroneis quadrata M. Peragallo & Heribaud Tempere & Peragallo 1909 

Stauroneis schinzii (Brun) Cleve Boyer 1927b 

Stauroneis semen Ehrenberg Stoermer & Kreis 1978 

Stauroneis siberica (Grunow) Lange-Bertalot Stoermer et al. 1999 

Stauroneis sieboldii Ehrenberg Patrick & Reimer 1966 

Stauroneis smithii Grunow Stoermer & Kreis 1978 

Stauroneis smithii var. borgei Ehrenberg Stoermer et al. 1999 

Stauroneis smithii var. incisa Pantocsek Cambum 1982 

Stauroneis smithii var. minima Haworth Stoermer & Kreis 1978 

Stauroneis staurolineata Reimer Patrick & Reimer 1966 

Stauroneis stodderi Greenleaf in Lewis Stoermer & Kreis 1978 

Stauroneis subula Ehrenberg 1856 

Stauroneis tenera Hustedt Hohn & Hellerman 1963 

Stauroneis terryi Ward in Palmer Boyer 1927b 

Stauroneis thermicola (Petersen) Lund Dodd 1987 

Stauroneis triundulatum Borge Patrick & Reimer 1966 

Stauroneis virginica Ehrenberg 1856 

Stauroneis wislouchii Poretzky & Anisimata Grimes & Rushforth 1982 

Staurophora amphioxys (Gregory) D.G Mann Stoermer et al. 1999 

Stauroptera aspera Ehrenberg 1856 

Stauroptera isostauron Ehrenberg 1856 

Stauroptera legumen Ehrenberg 1 856 

Stauroptera microstauron Ehrenberg 1856 

Stauroptera parva Ehrenberg 1 856 

Stauroptera trinodis Ehrenberg 1 856 

Staurosira brevistriata Grunow ~ Stoermer & Kreis 1978 

Staurosira capucina Borzsc Stoermer & Kreis 1978 

Staurosira construens Ehrenberg Williams & Round 1987 

Staurosira construens var. binodis (Ehrenberg) Hamilton in Hamilton et al Hamilton et al. 1992 

Staurosira construens var. capitata (Heribaud) Andresen et al Andresen et al. 2000 

Staurosira construens var. minuta (Tempere & Peragallo) Andresen et al Andresen et al. 2000 

Staurosira construens f. subsalina (Hustedt) Andresen et al Andresen et al. 2000 

Staurosira construens var. venter (Ehrenberg) Hamilton Stoermer et al. 1999 

Staurosira elliptica (Schumann) Wiliams & Round Morales 2001 

Staurosira entomon Ehrenberg Stoermer & Kreis 1978 

Staurosira leptostauron var. dubia (Grunow) Edlund Stoermer et al. 1999 

Staurosira mutabilis (W. Smith) Grunow Stoermer & Kreis 1978 

Staurosira pinnata Ehrenberg 1 856 



474 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Name Publication 

Staurosira stevensonii Manoylov. Morales & Stoermer Manoylov et al. 2003 

Staurosira venter (Ehrenberg) Grunow in Pantocsek Siver et al. 2005 

Staurosirella berolinensis (Lemmerman) Edlund Morales 2002 

Staurosirella lapponica (Grunow in Van Heurck) Williams & Round Stoermer et al. 1999 

Staurosirella leptostauron (Ehrenberg) Williams & Round Stoermer et al. 1999 

Staurosirella leptostauron var. dubia (Grunow) Bukhtiyarova Morales 2002 

Staurosirella leptostauron var. fossilis (Pantocsek) Andresen et al Andresen et al. 2000 

Staurosirella leptostauron var. rhomboides (Grunow) Andresen et al Andresen et al. 2000 

Staurosirella pinnata (Ehrenberg) Williams & Round Williams & Round 1987 

Staurosirella pinnata var. intercedens (Grunow in Van Heurck) Hamilton Stoermer et al. 1999 

Staurosirella pinnate var. lancettula (Schumann) Siver & Hamilton in Siver et al Siver et al. 2005 

Staurosirella trigona (Brun & Heribaud) Siver & Hamilton in Siver et al Siver et al. 2005 

Stenopterobia anceps (Lewis) Brebisson Stoermer et al. 1999 

Stenopterobia anceps f. subacuta Fricke Hustedt 1912 

Stenopterobia curvula (W. Smith) Krammer Dute et al. 2000 

Stenopterobia delicatissima (Lewis) Brebisson Dute et al. 2000 

Stenopterobia densestriata (Hustedt) Krammer Gaiser & Johansen 2000 

Stenopterobia intermedia (Lewis) Fricke Stoermer & Kreis 1978 

Stenopterobia intermedia f. densistriata Hustedt Whitford & Schumacher 1973 

Stenopterobia intermedia f. subacuta Fricke Cambum 1982 

Stenopterobia intermedia f. undulata Sovereign Sovereign 1963 

Stenopterobia rautenbachiae Patrick & Roberts 1979 

Stephanodiscus alpinus Hustedt Stoermer & Kreis 1978 

Stephanodiscus astraea (Ehrenberg) Grunow Stoermer & Kreis 1978 

Stephanodiscus astraea var. intermedia Fricke Stoermer & Kreis 1978 

Stephanodiscus astraea var. minutula Grunow Stoermer & Kreis 1978 

Stephanodiscus astraea var. minutulus (Kiitzing) Grunow Stoermer & Kreis 1978 

Stephanodiscus astraea var. spinulosa Tempere & Peragallo 1909 

Stephanodiscus barkleyi Rattray Whitford 1956 

Stephanodiscus binatus Hakansson & Kling Siver et al. 2005 

Stephanodiscus binderanus (Kiitzing) Kreiger Stoermer & Kreis 1978 

Stephanodiscus binderanus var. oestrupii (A. Cleve) A. Cleve Stoermer & Kreis 1978 

Stephanodiscus carconensis Grunow Stoermer & Kreis 1978 

Stephanodiscus carconensis var. minor Tempere & Peragallo 1912 

Stephanodiscus carconensis var. pusilla Grunow Stoermer & Kreis 1978 

Stephanodiscus conspicueporus Stoermer, Hakansson & Theriot Stoermer et al. 1999 

Stephanodiscus dubius (Fricke) Hustedt Stoermer & Kreis 1978 

Stephanodiscus excentricus Hustedt Kociolek & Herbst 1992 

Stephanodiscus hantzschii Grunow Stoermer & Kreis 1978 

Stephanodiscus hantzschii var. delicatula Cleve Whitford & Schumacher 1973 

Stephanodiscus hantzschii f. tenuis (Hustedt) Hakansson & Stoermer Stoermer et al. 1999 

Stephanodiscus hantzschii var. pusilla Grunow Stoermer & Kreis 1978 

Stephanodiscus invisitatus Hohn & Hellerman Stoermer & Kreis 1978 

Stephanodiscus kuetzingiana (nomen nudum) Stoermer & Kreis 1978 

Stephanodiscus lucens Hustedt Stoermer et al. 1999 

Stephanodiscus medius Hakansson Stoermer et al. 1999 

Stephanodiscus minutula (Kiitzing) Round Stoermer et al. 1999 

Stephanodiscus minutulus (Kiitzing) Cleve & MoUer Stoermer et al. 1999 

Stephanodiscus minutus Grunow Stoermer & Kreis 1978 

Stephanodiscus niagarae Ehrenberg Stoermer & Kreis 1978 

Stephanodiscus niagarae var. magniflca Fricke Stoermer & Kreis 1978 

Stephanodiscus parous Stoermer & Hakansson Stoermer et al. 1999 

Stephanodiscus reimerii Theriot & Stoermer Theriot 1992 

Stephanodiscus rotula (Kiitzing) Hendey Stoermer & Kreis 1978 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 475 



Name Publication 

Stephanodiscus rotula var. minutula (Kiitzing) Grunow Stoermer & Kreis 1978 

Stephanodiscus subsalsus (A. Cleve) Hustedt Stoermer & Kreis 1978 

Stephanodiscus subtilis (Van Goor) A. Cleve Stoermer & Kreis 1978 

Stephanodiscus subtransilvanicus Gasse Stoermer et al. 1999 

Stephanodiscus superiorensis Stoermer & Theriot Stoermer et al. 1999 

Stephanodiscus tenuis Hustedt Stoermer & Kreis 1978 

Stephanodiscus transylvanicus Pantocsek Stoermer & Kreis 1978 

Stephanodiscus vestibulis Hakansson, Stoermer & Theriot Stoermer et al. 1999 

Stephanodiscus yellowstonensis Theriot & Stoermer Therior & Stoermer 1984 

Stephanosira mississippica Ehrenberg 1856 

Striatella fenestrata (Lyngbye) Kuntze Stoermer & Kreis 1978 

Striatella flocculosa (Roth) Kuntze Stoermer & Kreis 1978 

Surirella acredula Hohn & Hellerman Hohn & Hellerman 1963 

Surirella adumbratus Hohn & Hellemian Hohn & Hellerman 1963 

Surirella agmatilis Cambum Cambum 1982 

Surirella alicula Hohn & Hellerman Collins & Kalinsky 1977 

Surirella amphioxys W. Smith Stoermer & Kreis 1978 

Surirella anceps Lewis Stoermer & Kreis 1978 

Surirella angusta Kutzing Stoermer & Kreis 1978 

Surirella apiculata W. Smith Stoermer & Kreis 1978 

Surirella arcta A. Schmidt Tempere & Peragallo 1909 

Surirella arctissima A. Schmidt Boyer 1927b 

Surirella baileyi Lewis Boyer 1927b 

Surirella barca Hohn & Hellerman Hohn & Hellerman 1963 

Surirella beadensis Sovereign Sovereign 1963 

Surirella bifrons Ehrenberg Stoermer & Kreis 1978 

Surirella birostrata Hustedt Stoermer & Kreis 1978 

Surirella biseriata Brebisson & Godey Stoermer & Kreis 1978 

Surirella biseriata var. bifrons (Ehrenberg) Hustedt Stoermer & Kreis 1978 

Surirella biseriata var. bifrons f. amphioxys (W. Smith) Hustedt Whitford & Schumacher 1973 

Surirella biseriata var. bifrons f. punctata Meister Stoermer et al. 1999 

Surirella biseriata var. constricta Grunow Whitford & Schumacher 1973 

Surirella biseriata var. diminuta A. Cleve Stoermer & Kreis 1978 

Surirella biseriata var. orientalis Skvortzow Stoermer et al. 1999 

Surirella biseriata var. subacuminata Grunow Clark & Rushforth 1977 

Surirella bohemica Maly Clark & Rushforth 1977 

Surirella brebissonii Krammer & Lange-Bertaiot Stoermer et al. 1999 

Surirella brebissonii var. kuetzingii Krammer & Lange-Bertalot Potapova & Charles 2003 

Surirella brightwellii W. Smith Cambum 1982 

Surirella capronii Kitton Boyer 1927b 

Surirella cardinalis Kitton Stoermer & Kreis 1978 

Surirella carolinensis Ehrenberg 1856 

Surirella carolinicola Cambum Cambum 1982 

Surirella circumsuta Ehrenberg 1 856 

Surirella cocconeis Ehrenberg 1856 

Surirella constricta Ehrenberg 1856 

Surirella craticula Ehrenberg Rushforth & Merkley 1988 

Surirella crenulata Ehrenberg H.L. Smith 1876-1888 (#517) 

Surirella cruciata A. Schmidt Tempere & Peragallo 1909 

Surirella crumena Brebisson in Kutzing Boyer 1927b 

Surirella cuspidata Hustedt Cambum & Charies 2000 

Surirella delicatissima Lewis Stoermer & Kreis 1978 

Surirella delicatissima f. tenuissima Manguin Dixit & Smol 1995 

Surirella didyma Kutzing Stoermer & Kreis 1978 



476 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Name Publication 

Surirella elegans Ehrenberg Stoermer & Kreis 1978 

Surirella elegans f. minor Tempere & Peragallo 1908 

Surirella elegans var. norwegica (Eulenstein) Brun Clark & Rushforth 1977 

Surirella elliptica Terry Tempere & Peragallo 1908 

Surirella engleri Patrick & Roberts 1979 

Surirella eximia Ehrenberg 1856 

Surirella fasciculata O. Miiller Stoermer et al. 1971 

Surirella flexuosa Ehrenberg 1856 

Surirella geroltii Ehrenberg H.L. Smith 1876-1888 (#517) 

Surirella gracilis (W. Smith) Grunow Stoermer & Kreis 1978 

Surirella gracilis var. gigantea Tempere & Peragallo Tempere & Peragallo 1909 

Surirella guatemalensis Ehrenberg Stoermer & Kreis 1978 

Surirella helvetica Brun Hansmann 1973 

Surirella intermedia Lewis Boyer 1927b 

Surirella iowensis Lowe Collins & Kalinsky 1977 

Surirella kittonii A. Schmidt Tempere & Peragallo 1908 

Surirella kittonii var. asperula M. Peragallo in Tempere & Peragallo Tempere & Peragallo 1908 

Surirella lagerheimii Cleve Stoermer & Kreis 1978 

Surirella librile Ehrenberg Stoermer & Kreis 1978 

Surirella limosa J.W. Bailey H.L. Smith 1876-1888 (#523) 

Surirella linea Ehrenberg 1 856 

Surirella linearis W. Smith Stoermer & Kreis 1978 

Surirella linearis var. constricta (Ehrenberg) Grunow Stoermer & Kreis 1978 

Surirella linearis var. helvetica (Brun) Meister Stoermer & Kreis 1978 

Surirella litoralis Hustedt Kalinsky 1983 

Surirella macra A. Schmidt Cleve & Moller 1879 

Surirella marylandica Ehrenberg 1856 

Surirella minuta Brebisson Stoermer & Kreis 1978 

Surirella molleriana Grunow Stoermer & Kreis 1978 

Surirella myodon Ehrenberg Ehrenberg 1 856 

Surirella nervosa A. Schmidt Aubert 1895 

Surirella nevadensis Hanna & Grant Kociolek & Herbst 1992 

Surirella norvegica Eulenstein Stoermer & Kreis 1978 

Surirella oblonga Ehrenberg Boyer 1927b 

Surirella oophaena Ehrenberg 1856 

Surirella oregonica Ehrenberg Stoermer & Kreis 1978 

Surirella oregonica f. minor Tempere & Peragallo Tempere & Peragallo 1909 

Surirella ovalis Brebisson Stoermer & Kreis 1978 

Surirella ovalis var. angusta (Kutzing) Van Heurck Tilden 1894-1909 (#248) 

Surirella ovalis var. baltica (Schumann) Cleve Benson & Rushforth 1975 

Surirella ovalis var. brightwellii (W. Smith) A. Cleve Grimes & Rushforth 1982 

Surirella ovalis var. minuta Van Heurck Stoermer & Kreis 1978 

Surirella ovata Kiitzing Stoermer & Kreis 1978 

Surirella ovata var. africana Cholnoky Cambum 1982 

Surirella ovata var. crumena (W. Smith) Hustedt Stoermer & Kreis 1978 

Surirella ovata var. pinnata (W. Smith) Rabenhorst Stoermer & Kreis 1978 

Surirella ovata var. salina (W. Smith) Rabenhorst Stoermer & Kreis 1978 

Surirella ovata var. subsalina Patrick 1968 

Surirella ovata var. utahensis Grunow Rushforth & Merkley 1988 

Surirella palmeri Boyer Boyer 1927b 

Surirella panduriformis W. Smith Stoermer & Kreis 1978 

Surirella parma Sovereign Sovereign 1963 

Surirella patella Ehrenberg Whitford & Schumacher 1973 

Surirella patella var. neupaueri (Pantocsek) Cleve-Euler Whitford & Schumacher 1973 

Surirella peisonis Pantocsek Stoermer et al. 1999 

Surirella pinnata W. Smith Boyer 1927b 

Surirella pinnata var. panduriformis (W. Smith) Hustedt Stoermer et al. 1999 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 477 



Name Publication 

Surirella plicata Ehrenberg 1 856 

Surirella pseudovalis Hustedt Grimes & Rushforth 1982 

Surirella pygmaea Ehrenberg 1 856 

Surirella rattrayi A. Schmidt Aubert 1 895 

Surirella regina Janisch in A. Schmidt Boyer 1927b 

Surirella regula Ehrenberg Ehrenberg 1 856 

Surirella robusta Ehrenberg Stoermer & Kreis 1978 

Surirella robusta var. armata Hustedt Stoermer et al. 1999 

Surirella robusta var. erosa Tempere & Peragallo 1 908 

Surirella robusta var. splendida (Ehrenberg) Van Heurck Stoermer & Kreis 1978 

Surirella robusta var. splendida f. hustedtiana (Mayer) Hustedt Bateman & Rushforth 1984 

Surirella robusta var. splendida f. punctata Hustedt Cambum 1982 

Surirella robustior MacKay Stoermer & Kreis 1978 

Surirella rudis Hustedt Czamecki et al. 1981 

Surirella saxonica Auerswald Stoermer & Kreis 1978 

Surirella sigmoidea Ehrenberg Stoermer & Kreis 1978 

Surirella spiralis Kutzing Stoermer & Kreis 1978 

Surirella splendida Ehrenberg Stoermer & Kreis 1978 

Surirella splendida var. nervosa A. Schmidt Tempere & Peragallo 1908 

Surirella splendida f. punctata Hustedt Sovereign 1958 

Surirella stalagma Hohn & Hellerman Cambum 1982 

Surirella stoermerii Lowe Lowe 1972-1973 

Surirella striata Leud.-Fortm Stoermer & Kreis 1978 

Surirella striatula Turpin Stoermer & Kreis 1978 

Surirella subsalsa W. Smith Collins & Kalinsky 1977 

Surirella suecica Grunow Hohn & Hellerman 1963 

Surirella suevica Zeller Stoermer & Kreis 1978 

Surirella tenera Gregory Stoermer & Kreis 1978 

Surirella tenera var. nervosa A. Schmidt Stoermer & &eis 1978 

Surirella tenera var. palmeri (Boyer) Hustedt Patrick 1945 

Surirella tenera var. robusta Tempere & Peragallo 1908 

Surirella tenera var. splendidula A. Schmidt Tempere & Peragallo 1908 

Surirella tenuis Mayer Stoermer et al. 1999 

Surirella tenuissima Hustedt Stoermer & Kreis 1978 

Surirella terrj'i Ward Boyer 1927b 

Surirella testudinella Ehrenberg 1856 

Surirella testudo Ehrenberg Ehrenberg 1856 

Surirella triumphans A. Schmidt Boyer 1927b 

Surirella turgida W. Smith Stoermer & Kreis 1978 

Surirella undata Ehrenberg Ehrenberg 1856 

Surirella undulata Ehrenberg Stoermer & Kreis 1978 

Surirella utahensis (Grunow) Hanna & Grant Rushforth & Merkley 1988 

Surirella valida Ehrenberg Tempere & Peragallo 1908 

Surirella valida var. erosa Tempere & Peragallo 1908 

Surirella valida var. triumphanus Tempere & Peragallo 1908 

Surirella verrucosa Pantocsek Hayak & Hulbary 1956 

Surirella virginica Ehrenberg 1 856 

Synedra actinastroides Lemmermann Stoermer & Kreis 1978 

Synedra acus Kutzing Stoermer & Kreis 1978 

Synedra acus var. angustissima Grunow Stoermer & Kreis 1978 

Synedra acus var. delicatissima (W. Smith) Van Heurck Patrick 1945 

Synedra acus var. radians (Kutzingj Hustedt Stoermer & Kreis 1978 

Synedra acuta Ehrenberg Patrick & Reimer 1966 

Synedra acutissimus Patrick & Reimer 1966 

Synedra aequalis Kutzing Boyer 1927a 

Synedra affinis Kutzing Stoermer & Kreis 1978 



478 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

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Name Publication 

Synedra affinis var. acuminata Grunow Rushforth & Merkley 1988 

Synedra affinis var. fasciculata (Kiitzing) Grunow Patrick & Reimer 1966 

Synedra affinis var. gracilis Grunow Collins & Kalinsky 1977 

Synedra affinis var. lancettula Grunow Rushforth & Merkley 1988 

Synedra amphicephala Kutzing Stoermer & JCreis 1978 

Synedra amphicephala var. asiatica Skvortzow Stoermer et al. 1999 

Synedra amphicephala var. austriaca (Grunow) Hustedt Stoermer & Kreis 1978 

Synedra amphicephala var. intermedia A. Cleve Stoermer et al. 1999 

Synedra amphioxys Ehrenberg 1856 

Synedra amphirhynchus Rabenhorst Boyer 1927a 

Synedra berolinensis Lemmermann Drum 1981 

Synedra biceps Kutzing Cambum 1982 

Synedra bicurvata Biene in Rabenhorst Boyer 1927a 

Synedra capensis Grunow Collins & Kahnsky 1977 

Synedra capitata Ehrenberg Stoermer & Kreis 1978 

Synedra chaseii Thomas Stoermer & Kreis 1978 

Synedra crotonensis Grunow Stoermer & Kreis 1978 

Synedra crotonensis Edwards Cleve & Moller 1878 

Synedra crotonensis var. prolongata Grunow Stoermer & Kreis 1978 

Synedra cyclopum Brutschy Stoermer & Kreis 1978 

Synedra cyclopum var. gibbosa Naegeli Prescott & Dillard 1979 

Synedra cyclopum var. robustum Schulz Patrick & Reimer 1966 

Synedra danica Kutzing Stoermer & Kreis 1978 

Synedra debilis H.L. Smith 1876-1888 (#690) 

Synedra delicatissima W. Smith Stoermer & Kreis 1978 

Synedra delicatissima var. angustissima Grunow Stoermer & Kreis 1978 

Synedra delicatissima f. longissima Cleve & Moller 128 

Synedra delicatissima var. mesoleia Tempere & Peragallo 1909 

Synedra demerarae Grunow Stoermer & Kreis 1978 

Synedra dicephala Patrick & Reimer 1966 

Synedra dorsiventralis Muller Patrick & Reimer 1966 

Synedra entomon Ehrenberg 1856 

Synedra famelica Kiitzing Stoermer & Kreis 1978 

Synedra fasciculata (Agardh) Kutzing Stoermer & Kreis 1978 

Synedra fasciculata var. truncata (Greville) Patrick Stoermer & Kreis 1978 

Synedra filiformis Grunow Stoermer & Kreis 1978 

Synedra filiformis var. exilis A. Cleve Stoermer & Kreis 1978 

Synedra flexuosa Ehrenberg 1856 

Synedra gaillonii (Bon,') Ehrenberrg Stoermer & Kreis 1978 

Synedra goulardi Brebisson Stoermer & Kreis 1978 

Synedra goulardi var. fluviatilis (Lemmermann) Frenguelli Stoermer et al. 1999 

Synedra homostriata Hohn Patrick & Reimer 1966 

Synedra hyalina Tempere & Peragallo Patrick & Reimer 1966 

Synedra hyperborea Grunow Stoermer & Kreis 1978 

Synedra hyperborea var. rostellata Grunow Stoermer & Kreis 1978 

Synedra incisa Boyer Stoermer & Kreis 1978 

Synedra laevigata Grunow Stoermer & Kreis 1978 

Synedra lanceolata Kutzing Stoermer & Kreis 1978 

Synedra longiceps Ehrenberg Stoermer & Kreis 1978 

Synedra longissima W. Smith Stoermer & Kreis 1978 

S\'nedra lunaris Ehrenberg Stoermer & Kreis 1978 

Synedra mazamaensis Sovereign Stoermer et al. 1999 

Synedra minuscula Grunow Stoermer & Kreis 1978 

Synedra montana Krasske Stoermer & Kreis 1978 

Synedra nana Meister Cambum 1982 

Synedra netronoides Hohn & Hellerman Collins & Kalinsky 1977 

Svnedra notha Hohn & Hellerman Patrick & Reimer 1966 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 479 



Name Publication 

Svnedra obtusa W. Smith Boyer 1 927a 

SyTiedra ostenfeldii (Krieger) A. Cleve Stoermer et al. 1999 

Svnedra oxyrhynchus Kutzing Boyer 1927a 

SjTiedra oxwhynchus var. undulata Grunow Patrick & Reimer 1966 

Svnedra parallelogram Patrick & Reimer 1966 

Synedra parasitica (W. Smith) Hustedt Cambum 1982 

Svnedra parasitica var. subconstricta (Grunow) Hustedt Cambum 1982 

Synedra par\Iua Kutzing Boyer 1927a 

Synedra pulchella Ralfs ex Kutzing Cambum 1982 

S\Tiedra pulchella var. abnormis Macchiati Boyer 1927a 

Synedra pulchella var. flexella Boyer Boyer 1927a 

Synedra pulchella var. lacerata Hustedt Cambum 1982 

Synedra pulchella var. lanceolata O'Meara Whitford & Schumacher 1973 

Synedra pulchella f. major Gmnow in Van Heurck Patrick & Reimer 1966 

Synedra reca\ a Hohn Patrick & Reimer 1966 

Synedra radians Kutzing Camburn 1982 

Synedra mmpens Kutzing Camburn 1982 

Synedra mmpens var. familiaris (Kutzing) Hustedt Cambum 1982 

Synedra mmpens var. fragilariodes Gmnow Cambum 1982 

Synedra mmpens var. meneghiniana Gmnow Camburn 1982 

Synedra mmpens var. scotica Gmnow Camburn 1982 

Synedra scalaris Ehrenberg Patrick & Reimer 1966 

Synedra simalongis W. Smith Stoermer et al. 1999 

Synedra socia Wallace Collins & Kalinsky 1977 

Synedra spathulifera Gmnow Boyer 1927a 

Synedra spectabilis Ehrenberg 1856 

Synedra splendens Kutzing Boyer 1927a 

Synedra stauroneis Ehrenberg 1856 

Synedra stela Hohn & Hellerman Stoermer et al. 1999 

Synedra subaequalis (Gmnow) Van Heurck Smith 1950 

Synedra subrhombica Nygaard Camburn & Charles 2000 

Synedra subtilis Kiitzing H.L. Smith 1876-1888 (#579) 

Synedra tabulata (Agardh) Kutzing Camburn 1982 

Synedra tabulata var. accuminata Gmnow Drum 1981 

Synedra tabulata var. obtusa (Amott) Cleve Kaczmarska & Rushforth 1983 

Synedra tenera W. Smith Collins & Kalinsky 1977 

Synedra tenera f. elongata Cleve & Moller 1878 

Synedra tenuissima Kutzing Stoermer et al. 1999 

Synedra ulna (Nitzsch) Ehrenberg Camburn 1982 

Synedra ulna var. aequalis (Kutzing) Hustedt Camburn 1982 

Synedra ulna var. amphirhynchus (Ehrenberg) Gmnow Camburn 1982 

Synedra ulna var. biceps (Kutzing) Kirchner Stoermer et al. 1999 

Synedra ulna var. capitata Ehrenberg Patrick & Reimer 1966 

Synedra ulna var. chaseana Thomas Stoermer et al. 1999 

Synedra ulna var. claviceps Venkataraman Stoermer et al. 1999 

Synedra ulna var. constricta Venkataraman Hohn 1961 

Synedra ulna var. contracta 0stmp Camburn 1982 

Synedra ulna var. danica (Kiitzing) Van Heurck Camburn 1982 

Synedra ulna var. delicatissima Patrick & Reimer 1966 

Synedra ulna var. impressa Hustedt Stoermer et al. 1999 

Synedra ulna var. impressa f. contracta Patrick 1968 

Synedra ulna var. lanceolata Gmnow Tempere & Peragallo 1908 

Synedra ulna var. longissima (W. Smith) Bmn Collins & Kalinsky 1977 

Synedra ulna var. obtusa Van Heurck Collins & Kalinsky 1977 

Synedra ulna var. oxyrhynchus (Kutzing) Van Heurck Cambum 1982 

Synedra ulna var. oxyrhynchus f. mediocontracta (Forti) Hustedt Whitford & Schumacher 1973 

Synedra ulna var. radians Patrick & Reimer 1966 



480 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 27 

Name Publication ^ 

Synedra ulna var. ramesi (Heribaud) Hustedt Cambum 1982 

Synedra ulna var. spathulifera Grunow Collins & Kalinsky 1977 

Synedra ulna var. splendens Kiitzing Collins & Kalinsky 1977 

Synedra ulna var. subaequalis (Grunow) Van Heurck Rushforth & Squires 1985 

Synedra ulna var. vitraea Van Heurck Tempere & Peragallo 1908 

Synedra utermohlii Hustedt Stoermer et al. 1999 

Synedra valens Ehrenberg Patrick & Reimer 1966 

Synedra vaucheriae Kutzing Boyer 1927a 

Synedra vaucheriae var. capitellata (Grunow) Cleve Stoermer et al. 1999 

Synedra vaucheriae var. truncata (Greville) Grunow Stoermer et al. 1999 

Synedra vitrea Kutzing Boyer 1927a 

Tabellaria binalis (Ehrenberg?) Grunow Boyer 1927a 

Tabellaria fenestrata (Lyngbye) Kutzing Cambum 1982 

Tabellaria fenestrata var. asterionelloides Grunow Stoermer et al. 1999 

Tabellaria fenestrata var. geniculata A. Cleve Stoermer et al. 1999 

Tabellaria fenestrata var. gracilis Meister Patrick & Reimer 1966 

Tabellaria fenestrata var. intermedia Grunow Stoermer et al. 1999 

Tabellaria flocculosa (Roth) Kutzing Cambum 1982 

Tabellaria flocculosa var. linearis Koppen Stoermer et al. 1999 

Tabellaria nodosa Ehrenberg 1856 

Tabellaria quadriseptata Knudson Stoermer et al. 1999 

Tabellaria silicula Ehrenberg 1856 

Tabellaria teilingii Patrick & Reimer 1966 

Tabellaria trinodis Ehrenberg Stoermer et al. 1999 

Tabellaria venter Ehrenberg Patrick & Reimer 1966 

Tabellaria ventricosa Kiitzing Aubert Le Diatomiste #20 

Tabellaria vulgaris Ehrenberg Patrick & Reimer 1966 

Tabularia fasciculata (C. Agardh) Williams & Round Stoermer et al. 1999 

Terpsinoe americana (Bailey) Ralfs Boyer 1927a 

Terpsinoe intermedia Grunow Kaczmarska & Rushforth 1983 

Terpsinoe musica Ehrenberg Stoermer & Kreis 1978 

Tetracyclus elliptica (Ehrenberg) Grunow Patrick & Reimer 1966 

Tetracyclus emarginatus (Ehrenberg) W. Smith Patrick & Reimer 1966 

Tetracyclus lacustris Ralfs Patrick & Reimer 1966 

Tetracyclus mpestris (A. Braun) Gmnow Stoermer & Kreis 1978 

Tetracyclus stella Ehrenberg Tempere & Peragallo 1909 

Tetracyclus rhombus var. maxima Tempere & Peragallo 1909 

Tetragramma americana Ehrenberg 1 856 

Thalassiocyclus lucens Hakansson & Mahood Stoermer et al. 1999 

Thalassiosira bramaputrae (Ehrenberg) Hakansson & Locker Stoermer et al. 1999 

Thalassiosira fluviatilis Hustedt Stoermer & Kreis 1978 

Thalassiosira guillardii Hasle Stoermer et al. 1999 

Thalassiosira incerta Makarova Stoermer et al. 1999 

Thalassiosira lacustris (Gmnow) Hasle Stoermer et al. 1999 

Thalassiosira levanderi Van Goor Stoermer & Kreis 1978 

Thalassiosira pseudonana Hasle & Heim Stoermer & Kreis 1978 

Thalassiosira simplex Hustedt Stoermer et al. 1999 

Thalassiosira visurgis Hustedt Stoermer & Kreis 1978 

Thalassiosira weissflogii (Gmnow) Fryxell & Hasle Cambum 1982 



KOCIOLEK: FRESHWATER DIATOMS OF INLAND ENVIRONMENTS IN THE U.S. 48 1 



Name Publication 

Triceratium jensenianum Grunow Collins & Kalinsky 1977 

Triceratium solenoceros Ehrenberg Collins & Kalinsky 1977 

Triceratium venosum Brighton Collins & Kalinsky 1977 

Tropidoneis lepidoptera (Gregory) Cleve Stoermer & Kreis 1978 

Tropidoneis lepidoptera var. proboscidea Cleve Stoermer & Kreis 1978 

Tropidoneis vitrea (W. Smith) Cleve Rushforth & Merkley 1988 

Tropidoneis vitrea var. scaligera (Grunow) Cleve Rushforth & Merkley 1988 

Trybhonella acuta (Cleve) D.G Mann Stoermer et al. 1999 

Tryblionella angustata W. Smith Stoermer et al. 1999 

Tryblionella angustata var. acuta (Grunow) Bukhtiyarova Stoermer et al. 1999 

Trybhonella apiculata Gregory Stoermer et al. 1999 

Trybhonella debihs (Amott) Grunow in Cleve & Grunow Stoermer et al. 1999 

Trybhonella gracilis W. Smith Stoermer et al. 1999 

Trybhoneha hungarica (Grunow) D.G Mann Stoermer et al. 1999 

Trybhoneha levidensis W. Smith Stoermer & Kreis 1978 

Trybhonella plana var. fennica (Hustedt) Simola Siver et al. 2005 

Tr>'blionella scutellum (Bailey) W. Smith Stoermer & Kreis 1978 

Trybhonella victoriae Grunow Stoermer et al. 1999 

Ulnaria ulna (Nitzsch) Compere Siver et al. 2005 

Urosolenia eriensis (H.L. Smith) Round & Crawford Stoermer et al. 1999 

Urosloenia gracilis (H.L. Smith) Andresen et al Andresen et al. 2000 

Urosolenia longiseta (Zach.) Edlund & Stoermer Stoermer et al. 1999 

Vanheurckia rhomboides (Ehrenberg) Brebisson Patrick 1945 

Vanheurckia rhomboides var. amphipleuroides (Brebisson) Van Heurck Patrick 1945 

Vanheurckia rhomboides var. crassinervia (Brebisson) Van Heurck Patrick 1945 

Vanheurckia rhomboides var. crassinervia f. capitata Patrick Patrick 1945 

Vanheurckia viridula Brebisson Tempere & Peragallo 1908 

Vanheurckia vulgaris (Thwaites) Van Heurck Patrick 1945 



Section III: Bibliography 



Abbott, M.R., Powell, T.M. and Richerson, RJ. 1982. The relationship of environmental variability to the 
spatial patterns of phytoplankton biomass in Lake Tahoe. Journal of Plankton Research 4:927-941. 

Ahlstrom, E.H. 1936. The deep-water plankton of Lake Michigan, exclusive of the Crustacea. Transactions 
of the American Microscopical Society 55:286-299. 

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Copyright © 2005 by the California Academy of Sciences 
San Francisco, California, U.S.A. 



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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 

Volume 56. No. 28. pp. 527-556. 29 figs., 2 tables. December 30, 2005 



Physical Geography of the GaoUgong Shan Area of 
Southwest China in Relation to Biodiversity 

George Chaplini 

Department of Anthropology, California Academy of Sciences, 875 Howard Street, 
San Francisco, California 94103; Email: gchaplin@ calacademy.org 

The Gaohgong Shan mountains (Gaoligong Shan) comprise the western-most part of 
the Hengduan Mountain Range. They include all of the contiguous ridges west of the 
Nujiang River and east of the Irrawadi-Nmai Rivers and lie at the junction of the 
Indo-Malaya and Palearctic zoogeographic realms. The Gaoligong Shan are one of 
the world's most significant biodiversity hotspots outside of the tropics. 

The Hengduan Mountains, of which the Gaoligong Shan are a part, are a result of 
the collision of the South China Block and Eurasian Plate during the late Mesozoic. 
During the Cenozoic, the Gaoligong Shan have also been affected by the continuing 
movements of the Indo-Australasian Plate and Eurasian Plates to the west of the 
Hengduan Mountains. 

The Gaoligong Shan are characterized by a number of unusual features. Their 
high, contiguous ridges extend further south than do most of those of the other 
Hengduan Mountains. Also, their river valleys are unusually narrow and deep 
because they are incised into hard rock that maintains steep slope profiles. 
Continuing uplift, steep gradients, and swiftly flowing rivers have eroded deep 
gorges. The north-south orientation of the river valleys causes the Gaoligong Shan to 
have an unusual face aspect relative to the sun; nearly all slopes face either east or 
west. The deep valleys and north-south orientation of the ridges result in the region 
having a more moderate climate than surrounding non-mountainous areas situated 
at the same latitude. Because of their antiquity, the Gaoligong Shan have accumulat- 
ed a high level of biodiversity. At the same time, their high elevations and deep 
gorges have acted as barriers to migration for most terrestrial organisms. Moreover, 
because of their unusual climate and many protected environments, the Gaoligong 
Shan provide a refugium from global climate perturbations. It is significant that the 
difficult terrain has, until recently, deterred extensive human habitation, thus pre- 
serving the region's biodiversity. 

Keywords: Gaoli Gongshan, Gaoligongshan, Hengduan Mountains, Biodiversity Hotspot, 

Climate, Refugia, Indian Plate, Australasian Plate, Eurasian Plate, Tibetan Plateau, 

GIS, Conservation, Biogeography. 

The Gaoligong Shan mountains (GLGS) are widely acknowledged as an important center of 
biodiversity and as such have been recognized as a World Heritage Site (UNESCO 2003). The list- 
ing recognizes the region's unusual geological context, ecological diversity, and scenic beauty. 
Although many authors have written about the GLGS, there is no general agreement as to the geo- 
graphical definition of the region. This paper presents such a definition. 

^ Research Associate, Department of Anthropology and Gaoligong Shan Informaticist, California Academy of 
Sciences . 

527 



528 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

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Ecologists recognize that the interplay of many factors is responsible for the accumulation of 
biodiversity. The prime factor is the environmental gradient produced by latitude and its effect on 
trophic production. The effect of latitude is modified by elevation and climatic variation due to 
local factors and ecological niche structure and complexity. The degree of isolation of the environ- 
ment controls interchange of species along the trophic levels. The age of the environment and its 
biotic components also promote biotic heterogeneity. Competition and predation interact with the 
niche structure to form biotic communities with various degrees of complexity. Generally, the older 
and more stable the area, the greater the biodiversity. Many aspects of mountain ecology serve to 
exaggerate the complexity of niche structure. Although many authors have alluded to different 
aspects of the ecology of the GLGS, none has reviewed them comprehensively in terms related to 
the physical geography. In this paper, the relationship of geography to ecologyis investigated with 
respect to the promotion and maintenance of biodiversity. 

In addition, the region's plate tectonics and geology are described. These two factors explain 
how the physical geography of the GLGS evolved. Tectonics causes rocks from different places to 
be brought together, building mountains or creating areas of subduction. The composition of the 
rocks dictates how they will behave as they are uplifted and eroded. Erosion of rocks contributes 
to soil formation, and the kind of soil formed is, in part, dependent on geology. The soils of the 
GLGS, therefore, are reviewed because of their bearing on biodiversity. The conformation of 
mountains and rivers has important biological consequences for the high levels of biodiversity in 
GLGS. 

General Considerations 

The purpose of this paper is to define the GLGS more accurately than has been done previous- 
ly. This requires a starting point. Broadly speaking, the GLGS. as referred to here, are the most 
westerly ridges of the Hengduan Mountains that extend north to south between the Nujiang River 
in the east and the Irrawadi River in the west. 

The name Gao-Li-Gong-Shan^ strictly speaking, applies to a single peak at the junction of 
Baoshan. Lushui. and Tengchong Counties at approximately 25.I33°N, 98.7 16°E. The exclusive 

^ In contracting Chinese names, I have used the established method Gao— Ift— Li— ^— Gong— '^— Shan— lL| 
of Zhao (1986) who suggested that names should not be contracted to less 

than two characters using the example of the Tian Shan Mountains, which he thought was preferable to Tian Mountains 
even though Shan means Mountains. Similarly, here I use Nujiang River rather than the Nu River despite the fact that 
"Jiang" means river. This follows the most common usage of the name in the non-Chinese literature. 

The name Gaoligong Shan in Chinese is complicated. It. like many Chinese names, has different layers of meaning. 
It can mean, literalh'. and based on the characters alone. High Multitude Tribute Mountain. The character Li that is used, is 
the same as the one used for the transliteration Li in the name of the people called Lisu and who are the dominant ethnic 
minority group of the area. The exclusive usage in Chinese is that Gaoligong Shan implies the whole range of which Mount 
Gaoligong Shan is in the middle. The alternati\e term that could be used is Shanmai, which means mountain range in 
Chinese. This would gi\ e the rather cumbersome and never used term Gaoligong Shanmai or more correctly, but even worse 
sounding. Gaoligong Shan Shanmai. In this paper, the name used for the whole range will also be Gaoligong Shan (GLGS); 
I recommend that alternative names, sometimes seen in print, should be avoided, e.g. "Gaoli's," "Gaoligong," "Gaoli 
Gongshan," or just "Gongshan." "Mount Gaoligong Shan" should be used for the single peak if needed. 

Another point: names in this region are difficult because of diverse ethnicities and dialects, e.g., the commonly 
referred to name Gongshan. is used sometimes for the mountains, at other times the county seat for the administrative zone, 
and most often, as an abbre\iation for an administrative zone itself. This is the Gongshan Dulong Nuzu Zizhixian. 
Sometimes the Gongshan administrative zone is referred to as the Dulong area. Officially, it should be the Gongshan 
Drungzu Nuzu Zizhixian (Carto. Pub. Hse 1984). and in this paper it will be referred to as (Gongshan County). Drung, 
Dulong, and even Delung are the name for same people but spelled in different dialects. The name used can have many 
implications (Gros 2004). In this paper, place names are given to be as informative as possible to the general reader (see 
also cartography section in the main text). Lastly. GLGS is used in the plural to signify the whole range, as we do for the 
Rocky Mountains of the western United States. 

Carto. Pub. Hse (1984). Map of the People's Republic of China. Cartographic Publishing House. Beijing: Esselte Map Ser\ice AB. Sweden. 
Gros, S. (2004). The Pohtics of Names: The Identification of the Dulong (Drung) of Northwest Yunnan. China Information. The 
Documentation and Research Centre for Modem China. Sinological Institute. Leiden University 18(2):275-302. 



CHAPLIN: GEOGRAPHY OF THE GAOLIGONG SHAN, SOUTHWEST CHINA 



529 








( V N 






fzogang r /\ 


r ) 


w 


( Zayu\x n^ 


( 

i 


X 

1 
) 


^ X SGongkrtan 

■* \ \Pu jong 
Kachin State j 1 1 

//^^«„^-.»,«Li Baoshan 
Y'lrmjiangyy,^ j/Xj^'^y 

lyy'^y-^^-^y^img 



FiGLiRE 2. Localor map showing the 
GaoHgong Shan highUghted in relation to 
local political geography. 



Figure 1 . Regional locator map showing the Gaoligong Shan highlight- 
ed in relation to political geography. 

usage in Chinese is that Gaoligong Shan includes the whole 
range of mountains of which the peak named Gaoligong Shan 
is in the middle. Chinese does not distinguish between plural 
and singular. 

The GLGS are a poorly known biodiversity hotspot in 
Southwest China located mostly in Yunnan Province. They 

are the most biodiversity-rich area (Lan and Dunbar 2000; Mackinnon et al. 1996) of Yunnan, 
which is China's most biodiverse province (Zhang and Lin 1985) (see Figs. 1 and 2). The GLGS 
occupy about 10.5% of Yunnan Province. 

The GLGS are a rugged mountainous border region adjoining Myanmar on the northeast. 
From a biogeographic perspective, the GLGS form the junction of several biogeographical realms, 
the Indo-Malayan, and Palearctic, and the biogeographical provinces of the Tibetan Plateau and 
South China subregion. The GLGS also stand at the junction of three major tectonic plates, which 
are discussed below, and, thus, 
three geological provinces. This 
position has given them an inter- 
esting and complex geological 
structure. 

The formation of the 
Hengduan Mountains, of which 
the GLGS are a part, preceded 
the uplift of the Qinghai-Xizang 
(Tibetan; Plateau. However, 
today the GLGS are, more or 
less, an extension of the Tibetan 
Plateau, which extends far to the 
south into Yunnan (see shaded 
area in Fig. 3). Subsequent to the 
uplift of the Plateau the Figure 3. Regional locator map showing the Gaoligong Shan highlighted 

. ' in yellow to the south of the Tibetan Plateau and to the west of the rest of the 

Hengduan Mountains and GLGS Hengduan Mountains. 




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Fourth Series. Volume 56, No. 28 




Figure 4. Satellite Image of Hengduan Mountains centered on Mt Gaoligong Shan, the Nujiang River can be seen enter- 
ing the snow covered Tibetan Plateau at the top of the image (NASA 2004a). 



experienced uplift, which has continued throughout much of the Cenozoic, and is associated with 
the Himalayan orogeny. The region is divided by a few large north-south flowing rivers, which are 
of major importance to Southeast Asia. The rivers run in extremely deep, gorges, which, having cut 
into the uplifting mountainous area, gave rise to a series of narrow, north-south-oriented, high 
mountain ridges. The rivers, which are associated with major fault zones (see Fig. 4), divide the 
area biogeographically. 

Tectonics 

The north-south orientation of the Hengduan Mountains is orthogonal to the predominant east- 
west mountains found throughout eastern Eurasia. The Chinese name "Hengduan" translates as the 
"Transverse" or "Transecting" Mountains. The GLGS comprise the most westerly mountain ridge 
of the Hengduan Mountains. 

Early Tectonic Movements. — The Hengduan Mountains were formed by several differ- 
ent major tectonic events. The mountains are at the margins of several plates, the Eurasian Plate to 
the north, the Indochina Block to the south, and the Indian Plate to the west. These plates are con- 
strained by the Philippine-Pacific Plates to the east, and the Australasian Plate to the south (Hall 
1997). These plates are all moving relative to the stable Eurasian Plate. The Hengduan Mountains 
region, being at the plate margins, is an active earthquake zone. 



CHAPLIN: GEOGRAPHY OF THE GAOLIGONG SHAN. SOUTHWEST CHINA 53 1 



The very earliest collision involved the subduction of plate fragments (blocks or terranes), 
including the Southern China Block, which were driven north and eastwards by the 
Philippine/Pacific Plates after they broke away from Pangea and Gondwana. These older move- 
ments underlie the eastern part of the Hengduan Mountains in northern Sichuan and their extension 
along the Longmen Mountains. From the initial breakup of Pangea, in the early Carboniferous 
between 350 to 300 Mya. the South China Block was always slightly ahead of the Indochina Block 
as both moved northwards. Nonetheless, the two were always closely associated. The Southern 
China Block contacted the Eurasian Plate shortly after the Northern China Plate and before the 
Indochina Block: this happened as early as 200 Mya. 

The Southern Terranes separated from the Northern Ten-anes very early, around 300 Mya. The 
sub-plate of the Southern Terranes broke away from Pangea much later than did the previously 
mentioned Plates and Blocks (South China, Indochina, and Northern Terranes). The Northern 
Terranes impacted the Eurasian Plate to the northwest of the South China Block. The Lhasa Terrane 
forms part of the GLGS and it belongs to the Northern Terrane group. The Southern Terranes man- 
aged to catch up with the Lhasa and Northern Terranes as they were slowed by collision into the 
Eurasian Plate. By 100 Mya, the Southern Terranes were abutting the Northern Terranes, on the 
south side of the Eurasian Plate and were adjoining the west side of the Hengduan Mountains. 
These plates can be seen in Figure 5 at 65 Mya and are already in place at that time. Figures 5-7 
were produced using the web service of the Ocean Drilling Stratigraphic Network Plate Tectonic 
Reconstruction Service (Soeding 2004). and the general discussion follows maps produced at the 
Histor}' of Global Plate Motions (Dutch 1998, citing data from Scotese 1994). The Hengduan 
Mountains are, thus, the result of these collisions. Formerly, the upper GLGS region would have 
been influenced by these early tectonic movements and collisions. However, in the GLGS, any 
early signal has now mostly been overwritten by subsequent events. 

Cenozoic Tectonic Movements. — In the early Tertiary, by 55 Mya, both Northern and 
Southern terrane groups were enclosed by the Indian Plate to the south, the South China Block to 
the east, and the Eurasian Plate to the north. The closing of the Tethyian seaway north of the GLGS 
was achieved by the relatively fast collision of the Indian Plate with the Eurasian Plate in the early 
Cenozoic. Eventually, both Northern and Southern terranes were sandwiched between the Indian 
Plate and the Eurasian Plate and became extruded and highly deformed (see Fig. 6 at 25 Mya). The 
present strains on these plates change orientation in the region of the northern GLGS from north- 
south to east-west (Bi 2004). 

The Indian Plate moves north by as much as 50 mm per year, and the area has absorbed some 
1500 km of deformation since first contact between India and Eurasia (Replumaz et al. 2004). In 
Myanmar, the amount of annual movement is only 35 mm (Socquet and Pubellier 2003). The 
crustal thickness over the Tibetan Plateau is much greater than elsewhere. Remnants of ocean crust 
lie now just west of the GLGS in Assam and can be seen as the white patch in Figure 7. The ter- 
ranes can be seen as thin strips sandwiched in between the larger plates. The congregation of plates 
in the region of the Hengduan Mountains can clearly be seen in Figure 7. 

As a consequence of the Indian Plate's collision into the Eurasian Plate, the eastern Himalaya 
syntaxis rotated clockwise, and crustal fragments of the Northern and Southern Terranes extruded 
southeastward. The extrusion was along the NW-trending Karakoram-Jiali, N-trending Gaoligong 
Shan, and Sagaing Faults (Lin et al. 2004). Ages of the faults indicate that deformation may have 
started from the south along the Sagaing Fault in Indochina and propagated toward the north along 
the Gaoligong Shan Fault. Subsequently, the deformation proceeded toward the northwest along 
the Jiali Fault and then the Karakoram Fault in southern Tibet. Such a deformation trend reflects 
continuous deformation caused by the northward indentation of the Indian Plate into the Eurasian 



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Fourth Series, Volume 56, No. 28 




Figures 5-7. (5): Reconstruction of Plate movements for 65 Mya; (6): at 25 Mya; (7): present. Present coastline is 
shown in red on the grey plates. White indicates sea floor. Images courtesy of Soeding (2004). 



Plate, which has continued during the whole of the Tertiary (Lin et al. 2004). 

The Hengduan Mountains are bound by a series of north-and-northwest-striking Cenozoic 
faults: to the west by the Gaoligong Shan and Batang-Lijiang strike-slip systems, to the east by the 
Longmen Mountain thrust belt and the Xiaojiang Fault, and to the south by the Red River fault 
shear zone (Wang et al. 2001). The Cenozoic deformational history of the eastern Indo-Asian col- 
lision zone may be divided into three stages: (1) Eocene-Oligocene (40-24 Myr) transpression in 
eastern Tibet starting in the Red River Shear Zone just below the GLGS; (2) early-middle Miocene 
(24-17 Myr) transtension in eastern Tibet: and (3) late Neogene-Quatemary east-west extension, 
widespread in eastern Tibet and Indochina, which created small basins to the east, west and south 
of the GLGS (Wang et al. 2001). 

QUATERN.\RY TECTONIC MOVEMENTS. — The newest tectonic arrival, the Australasian Plate, 



CHAPLIN: GEOGRAPHY OF THE GAOLIGONG SHAN, SOUTHWEST CHINA 533 



has driven the Indochina Block northwai-ds, crushing and distorting the latter 's northern front. The 
Indochina Block is highly deformed in the north but behaves approximately like a rigid block in 
the south (Wang et al. 2001 ). Secondary thrusting in the area is now active in the south of the GLGS 
as a result of the Australasian Plate subducting beneath Indonesia. For example, in Pupiao — a basin 
adjoining the GLGS along the east bank of the Nujiang River in Baoshan County — the 
Miocene/Pliocene soft coal beds are upUfted 70 m and tilted so that the adit entrance is at 60 
degrees (pers. obs.). 

The latest area to deform is south of the GLGS at the point where the Nujiang River first heads 
east, then south, and then southwest. The river courses of the Nanding River to the south of the 
study area, the Dayang, Wanding, and Longchuan Rivers run along associated en echelon faults in 
a newly established rupture zone. In addition, the Nujiang River is strongly diverted westwards by 
them. This zone of active faults dates from the Early Pleistocene. The ENE-WSW trending 
Longling-Lancang fault zone cuts across the earlier tectonics during the later period (Guo et al. 
2000). This extensional drag possibly resulted from the orthogonal friction of the Indian Plate mov- 
ing north along the middle of Myanmar (Socquet and Pubellier 2003) or from the Australasian 
Plate's impact along Sumatra and the Andaman Islands. The Tengchong region also exhibits a 
series of N-S faults, which contain the upper reaches of the Daying, Longchuan and Mingguan 
Rivers. 

Tectonics and Mountain Building in the Gaoligong Shan. — The Paleozoic and 
Mesozoic Era plate movements resulted in faulting, folding, and the formation of metamorphic 
rocks and magmatite. These tectonic features were formed in the Paleozoic Era with the breakup 
of Pangea. The same trends continued with added impetus throughout the Mesozoic as the plates 
assembled on the south side of the Eurasian Plate. Then, in the Cenozoic, the extensive regional 
fault system was activated as a result of the collision of the Indian Plate with the Eurasian Plate, 
and a collage of terranes and other plate fragments. The complex tectonics of the GLGS region has 
resulted in extensive orogeny and erosion. It has also resulted in volcanism, extensive metamor- 
phism. and local eruptions. And, in the GLGS, it has resulted in the uplift and exposure of rocks of 
much older periods. The movements of the tectonic plates and reentrant terranes were facilitated 
through a series of large strike-slip faults, as mentioned above. The courses of the Nujiang River, 
Lancang River, the northern part of the Jingsha River, and the Red River to the south, flow in these 
very large fault structures. The main branch of the Irrawadi River follows in the the course of 
another set of faults zones further to the west. The rivers have been entrained by the uplift, but, 
because of their huge watersheds, their large flows were sufficient to keep pace with the uplift 
through their down-cutting action. The Nujiang River Gorge Fault facilitates some 17 mm of slip 
per year along the Yunnan River Valley fault system (Socquet and Pubellier 2003), a fact highly 
relevant to the dams planned for the area. The constitution, trending, and formation period of the 
compressional. north-south older tectonics are totally different from those of east- west extension- 
al tectonic active in the Longling-Lancang Rupture Zone that formed in the Pleistocene. 

Earthquakes. — As a result of extensive and ongoing tectonic activity in the Hengduan 
Mountains, it is an active fault zone with many earthquakes (Meyerhoff et al. 1991). To the east of 
the Hengduan Mountains, along a line from the Longmen Mountains to east of the Lancang River, 
is an active zone of large earthquakes that have registered eight and above on the Richter Scale. 
The Nujiang River Fault, Lancang Fault, and the Red River Rupture Zone, are strike-slip faults that 
register earthquake magnitudes of typically less than eight and rarely above seven. In contrast, the 
Yarlung Zangbo (Brahmaputra) Fault Zone, which encompasses a system of low-angle thrust 
faults, experiences larger earthquakes, ones that often register over eight. Strike-slip faults rupture 
at lower magnitudes than do thrust faults, which are usually associated with subduction boundaries. 



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Fourth Series, Volume 56, No. 28 

At first, the axis of the zone of thrust faults seems to have been in Longmen Mountains but moved 
west to the Nujiang River in the Permian and now is along the Yarlung Zangbo fault system 
(Meyerhoff et al. 1991). Currently, the main thrusting activity has moved south from the Yarlung 
Zangbo to the Frontal and Main Boundary Thrust systems in northern India and Nepal. Large earth- 
quakes have the potential for tectonic damming of the rivers, that is major slides that often result 
from earthquakes generated by fault movement. For example, this happened on the Yi'ong Zangbo 
River just northwest of the GLGS. There a 33-km-2 lake formed behind a 2500 m by 60 m high 
dam in 2000. The dam subsequently failed, which resulted a catastrophic flood of over 100 km in 
length. Evidence of such damming, and scouring floods should be visible in the river terraces if 
they have occurred on the Nujiang River. 

Biodiversity Implications of Tectonics. — The paleo-separation and subsequent reaggre- 
gation of plates from Gondwana and their eventual collision with the Eurasian Plate brought 
diverse biotas together from different paleo-continents. The area's complex uplift history has fos- 
tered greater genetic diversity in the region because of complex patterns of exchange, isolation, 
adaptation, extinctions, and speciation. Of particular importance has been geographic division due 
to the tectonically-driven incision of the landmass by massive rivers that has given rise to oppor- 
tunities for vicariant events leading to further diversification. Tectonic activity has implications for 
the evolution of diverse host rock and soil types (see below). 

Geology 

Geological Province Overview. — The geologic provinces of the GLGS broadly agree 
with the boundaries of their tectonic elements. The GLGS contain three geological provinces: (1) 
the Lhasa Terrane from the Northern Terrane Group, which extends from the north along the 
Nujiang River valley to 70 kms south of Fugong Town; (2) the Himalaya Block of the Southern 
Terrane Group in the northwest near the Dulong River and south to near Gongshan Town; and (3) 
the Tenasserim-Shan Block of the Indochina Block, which includes all of the middle and southern 
GLGS. The Qintang Terrane forms the eastern border to the GLGS, but it is seen within only a tiny 
portion of the study area near Lishadi Village just north of Fugong Town, (see Figs. 2, 8). Each 
province has a set of geologic characteristics that distinguishes it from surrounding provinces. 
These characteristics may include the predominant lithologies, the age of the strata, and the struc- 
tural style (Steinshouer et al. 1997; Wandrey and Law 1997). 

Age of the Rocks of the Gaoligong Shan. — The Paleozoic Era GLGS formations are 
dominated by fault-, fold-, metamorphic-, and magmatite-deformed rocks. The major outcropping 
of Mesozoic Era rocks is more to the east of the study area along the Qintang Terrane; the rocks 
have been uplifted and folded, accompanied by compressional foreshortening, giving rise to the 
Nushan Mountains. However, smaller outcrops of Mesozoic rocks occur at both ends of the study 
zone (Fig. 9). To the southeast and generally along the western edge are large areas of Precambrian- 
age metamorphic rocks. Further to the west in the modern Burmese Basin (Myanmar), the 
Precambrian is overlain by Tertiary and Quateman,' sediments. Many sedimentar}' strata in the 
GLGS have been lifted to being nearly vertical. The whole southern Hengduan Mountains area 
underwent more folding and uplift throughout the Cenozoic Era. This high degree of folding led in 
the southern GLGS to the exposure of older Lower Paleozoic sequences, some as early as 
Cambrian. Cenozoic rocks in the GLGS include further metamorphic changes to host rocks and, 
locally, volcanism around Tengchong Town, in the southwest and west of the GLGS in Myanmar 
probably reflect extensional tectonics as a result of ongoing subduction of the eastern limb of the 
Indian plate. Both Tertiary and Quaternary volcanics and sediments have formed in Tengchong 



CHAPLIN: GEOGRAPHY OF THE GAOLIGONG SHAN, SOUTHWEST CHINA 



535 



County. During the Cenozoic. a series of exten- 
sional basins formed and can be seen as small 
patches around the GLGS in Figure 9. These 
basins are associated with the change in thrust 
direction in the Plio-Pleistocene. Recent geo- 
logical deposits consist of considerable scree 
and colluvium. alluvium, flood facies, and river 
terracing that can be seen in a few places in the 
river valleys. In the extreme north of the 
GLGS. there are extensive glaciers and paleo- 
glacial features. The glaciers are shrinking at an 
astonishing rate, as can be seen when compar- 
ing recent satellite photographs with photos 
taken in the 1970s: this is probably as a result 
of global warming. 

Generalized Geology 

Starting from the north, the geology of the 
GLGS will be examined in more detail and 
briefly discussed, because geology has an 
impact on present landforms and implications 
for biodiversity. The data are taken from two 
USGS open-file reports for Far East Asia and 
South Asia, respectively (Steinshouer et al. 
1997: Wandrey and Law 1997). The maps pre- 
sented in this paper that were derived from 
these data are not accurate beyond 1 km and the 
discussion is of regional overview or general- 
ized geologies only. 

Lhasa Terrane. — The Lhasa Terrane 
comprises the north of the GLGS area, in east- 
em Chayu County (Zayii Xian) and the south- 
em part of Zougong County (Zogang Xian) of 
the Tibetan Autonomous Region (Xizang 
Zizhiquj. The Lhasa Terrane also forms the 
northwestern part of Gongshan Dulong-Nu 
Autonomous County of Yunnan Province 
(Gongshan Dmngzu-Nuzu Zizhixian) (referred 
to herewith as Gongshan County). The Lhasa 
Terrane is formed into a high mountainous area 
of Upper Paleozoic Rocks (PZu) (Figure 10). 
The Upper Paleozoic Rocks in general within 
the GLGS consist of intercalated beds of car- 
bonate, argillaceous deposits, basalts, and 
metamorphosed rocks with the upper facies 
containing more volcanics. The Lhasa Terrane 
is flanked to the northwest (north of the Dulong 




Figure 8. Map of the Geological Provinces within the 
Gaoligong Shan; these provinces broadly agree with the 
positions of tectonic plates (data from Steinshouer et al. 
1997. Wandrey and Law 1997). 




Figure 9. Geologic Map of the Gaoligong Shan showing 
the geologic age of the strata. 



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JK 
CF 



River) by Jurassic- Cretaceous (JK) age 
sequences. These sequences occur to the east 
where beds of Jurassic-Cretaceous age form the 
course of the Nujiang River. In the finger of the 
terrane extruded towards the town of Fugong, 
the river cuts through the Triassic (Tr) beds of 
the Lhasa or Qintang Terrane. Beyond, the 
Nujiang River reaches the Precambrian (pC) 
beds, which consist of some paratethys and 
some metamorphic and basaltic rocks. These 
Precambrian rocks form most of the ridge of 
the GLGS. The Precambrian metamorphics 
also form the ridges in the extreme west of 
Gongshan County and part of the ridge of the 
Patkai Range. The Patkai Range and other 
ridges in Myanmar are not part of the GLGS 
and lie outside of the study zone. Small patch- 
es of these Precambrian rocks are also exposed 
at the junction of the Himalaya Block, Lhasa 
Terrane, and the Tenasserim-Shan Block near 
to the town of Gongshan. 

Himalayan Block. — The Himalayan 
Block forms the southwest of Zayli County and 
forms all of the eastern part of Gongshan 
County west of the Dulong River watershed. 
The mountains of the Himalaya Block are 
lower than those of the Lhasa Terrane. The 
Himalayan Block is formed mostly of 
Mesozoic intrusive and metamorphic rocks 
(Mzim). The Himalayan Block also extends 
across the northern end of the Tenasserim-Shan 
Block to form the finger that is caught between 
the Tenasserim-Shan Block and the Lhasa 
Teirane. This finger is formed of Triassic meta- 
moiphic and sedimentary rocks (Trms), possi- 
bly a shallow sea ophiolitic melange. To the 
west, mostly outside of the study area, in the 
northwest corner of Gongshan County, there 
are extensive areas of Precambrian (pC) rocks 
belonging to the Himalaya Block. Between 
these and the metamorphic core of the 
Himalaya Block in Gongshan County is a flank 

of Carboniferous sedimentary rocks (Cs). On the other flank, between the westerly edge of the 
Himalaya Block and the Lhasa Terrane, are small outcroppings of Permian sedimentary rocks, 
probably consisting of deep-water turbidites (Meyerhoff et al. 1991). 

Tenasserim-Sr\n Block. — Main Ridge of the Gaoligong Shan in Fugong, Lushui, 
AND Kachin. — The majority of the GLGS sits on the Tenasserim-Shan Block. The middle reach- 




FlGURE 10. 

Gaoligong Shan 



s 

geologic units 



CHAPLIN: GEOGRAPHY OF THE GAOLIGONG SHAN, SOUTHWEST CHINA 537 



es of the GLGS comprise Fugong and Lushui Counties and to the west, Kachin State of Myanmar. 
The middle reaches of the GLGS ridge are in the northern part of the Tenasserim-Shan Block that 
extends south from the border with Gongshan County and the Himalaya Block. The GLGS main 
ridgeline skirts around the northern edge of Triassic metamorphic and sedimentary rocks (Trms) 
and runs south through the middle reaches between Precambrian (pC) on the east and Permian (Pr) 
and more Triassic metamorphic and sedimentary rocks (Trms). The Permian beds have more 
Teth}'ian affinities. To the west, the middle reaches of the GLGS are formed of Upper Paleozoic 
rocks (PZU). The actual ridge and the eastern flank of the GLGS are formed of Precambrian (pC) 
rocks. Small Carboniferous sedimentary (Cs) bodies pop out in the southern part of the middle sec- 
tion together with a larger Ordovician sedimentary outcrop, mostly grapholitic shales and meta- 
morphic rocks (Osm), probably composed of flysch and paraflysch. At the southern end of the mid- 
dle section, in Lushui County, occurs the end of the highest peaks where the ridgeUne is above 
3500m. 

Tenasserim-Shan Block: Main Ridge of the Gaoligong Shan in Baoshan 
Prefecture. — In the whole of the southern part of the GLGS, the main ridgeline is in Baoshan 
Prefecture. To the east of the main Ridgeline, the rocks are mostly Precambrian (pC). There are also 
outcrops of undifferentiated Paleozoic age of in northern Baoshan Prefecture; south of these are 
rocks of Carboniferous and Permian (CP) age. Around Daxue Mountain in Longling County in the 
south of Baoshan Prefecture, there are Lower Paleozoic rocks (PZl). At the eastern foot of 
Daxueshan at the southern extremity of the GLGS main ridgeline, in southeastern Longling 
County, is a Jurassic (Jr) intrusion. The Nujiang River flows east of this in a Silurian and 
Ordovician (SO) region between Longling and Shidian Counties. The western slope from the main 
GLGS ridge in Tengchong County down to the height of the Tengchong Basin is again Precambrian 
(pC) south to the Longchuan River. To the south of the Longchuan River in Longling County, the 
western and southern slopes of the main GLGS ridge are Lower Paleozoic rocks (PZl). 

Tenasserim-Shan Block: "NE-SW Trending Ridges" of Myanmar, and the counties 
OF Tengchong and Longjiang. — Because these ridges do not have a collective name, hereafter 
I will refer to them as the "NE-SW Trending Ridges." Between Lushui and Tengchong Counties, 
the border of China moves away from the GLGS main ridgeline into Myanmar along a line of high 
"NE-SW Trending Ridges." These ridges extend from Lushui County Line near Lushui Town 
towards Jiangao Mountain and continue further for 140 kms into Yingjiang County. The tectonic 
influences in the southern part and to the west are considerably younger, and have a tighter fold 
and fault structure imposed on the area. In the Myanmar part of the GLGS, there has been less 
uplift. Below Lushui County, the rock types become numerous, with smaller outcroppings. In the 
southern part of the GLGS south of Lushui County, there are similar sequence motifs between the 
western branch of "NE-SW Trending Ridges" and the eastern branch of the GLGS main mountain 
ridge, although the rock sequences are not identical. 

The westernmost flank of the "NE-SW Trending Ridges" is composed of rocks of Precambrian 
(pC) age. North of Tengchong County in Myanmar are Ordovician sedimentary and metamorphic 
(Osm) rocks, and southwest of these are more Carboniferous sedimentary (Cs) rocks. The 
Precambrian rocks return to be replaced further southwest by a small body of Cambrian (Cmsm) 
sediments and metasediments. Next to these Cambrian beds are more Ordovician sedimentary and 
metamorphic (Osm) rocks. The Precambrian makes up the south-westernmost corner of the GLGS 
ridges, except for inclusions of various igneous rocks. These parts of the "NE-SW Trending 
Ridges" are all in the Myanmar part of the GLGS. 

The eastern flank of the "NE-SW Trending Ridges " in northern Tengchong County is 
Ordovician/Silurian (OS) and Permian (Pr). In western Tengchong and Yingjiang Counties, the 



538 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 28 

eastern slopes of the "NE-SW Trending Ridges" are composed of Jurassic, Cretaceous, Triassic and 
other undifferentiated Mesozoic igneous rocks. West of these are small outcroppings of Tertiary 
volcanics, namely basaltic flows, andesitic lavas and pyroclastics, which overlie the Precambrian 
(pC) rocks that form the ridges of the "NE-SW Trending Ridges" in Myanmar. Just south of this, 
and slightly to the east in Yingjiang County, is an outcrop of Triassic (Tr) rocks along the border 
between Yunnan and Myanmar and the east flank of the "NE-SW Trending Ridges." 

Tenasserim-Shan Block: Central Basin Area of Yingjiang, Longchuan, and 
Tengchong Counties. — The area between the eastern flank of the "NE-SW Trending Ridges" 
and the western flank of the main ridgeline of the GLGS is a raised area dissected by the Daying 
and Longchuan Rivers and their tributaries. Running NE-SW down the middle of the area is a spine 
of Ordovician Silurian rocks that separates the drainage of the two rivers. There is a series of N-S 
to NE-SW trending faults that split the central area of Tengchong County from the "NE-SW 
Trending Ridges" down to the Longchuan River. These open up the Tengchong Basin into a "fan - 
folded" series of mountainous ridges. The upper headwaters of the Dayang and Longchuan Rivers 
and their tributaries like the Mingguan River run along these fault lines. These can be best seen in 
Figure 10. 

To the east of the Triassic igneous rocks, which occur along the border with Myanmar, there 
are Cambrian (Cm) and Silurian and Ordovician (SO) rocks, which extend towards the Longchuan 
River. To the east of the Longchuan River and north to the region of Tengchong Town, the area is 
mostly filled with Mesozoic beds of Jurassic-Cretaceous (JK) age, although these are extensively 
overlain by Neogene and Quaternary volcanic deposits and some younger volcanically-derived 
sediments. Tengchong County is characterized by a horseshoe-shaped opening to the south com- 
posed of Neogene sediments surrounding the Jurassic-Cretaceous mountains. The central region of 
Tengchong and eastern Yingjiang Counties is probably a zone of extension along the en echelon 
fault system that extends from here and further south. These younger faults cut across the older tec- 
tonic imprint (Guo et al. 2000). This extension would have exposed different rocks, as well as hav- 
ing allowed infilling by volcanic activity and for sedimentation to have taken place. These form the 
Quaternary Tengchong Basin, which appears to have subsequently uplifted. The Tengchong 
County pyroclastic cone field is not shown on Figure 10. It covers 600 km-2 and has erupted in five 
phases since the Tertiary. The nature of the eruption has changed from andesitic lavas in the early 
Tertiary to olivine-rich basalt lavas during the Pliocene through to the present. Daying Mountain 
Crater, 2865 m at 25.32°N, 98.47°E, last erupted in 1609 in an explosive eruption (Smithsonian 
2005). The many preserved cones in this area could be a source of local adaptation and vicariant 
speciation of smaller organisms. 

Soils 

The heterogeneity of soils in the GLGS is a consequence of the region's geologic diversity. 
The variety of host rocks, of very different ages, has given rise to many soil types. Although the 
remotely sensed data for this region are rather coarse ( 1 :4 M), some different dominant soil types 
are observable (F.A.O. 2005). The existence of many host rocks with different suites of predomi- 
nant minerals and a multitude of microclimates ensure a much greater diversity of soils on the 
ground than has been actually mapped. Within the study area, there exists a variety of soils with 
pH that varies from limestone-derived alkali types to acidic ones. 

In the north, on the Himalaya Block, are found the following lithosoils. humic cambisol, and 
eutric cambisol west of the Dulong. From the Lhasa Terrane in the north along the entire eastern 
slope and ridge of the GLGS are calcaric fluvisols. The western slope from the northern to the 
northwestern part of Tengchong County has lithosoil and humic cambisol. These continue south 



CHAPLIN: GEOGRAPHY OF THE GAOLIGONG SHAN. SOUTHWEST CHINA 539 



from Tengchong County along the western slope of the main GLGS ridge. In southwestern 
Tengchong and northern Yin^iang Counties are orthic acrisols. A tongue between them, from the 
central part of Tengchong County south until the Longling County border, is composed of ferric 
acrisol. The very southern portion of the study area in Longling County exhibits a more developed 
orthic acrisol (F.A.O. 2005). The predominant agricultural soil types seem to be latosols, laterite, 
red earths, yellow earths, purple earth, and paddy soils (F.A.O. 2005). These are mostly alluvial ter- 
races derived from the material of calcareous sedimentary rock, although the orthic acrisols are 
more acidic (RA.O. 2005). 

Soils evolve according to the latitude, elevation, temperature, and rainfall regime in which 
they are distributed. The same host rock minerals will give rise to different soils depending on the 
environment. Soil diversity, in turn, gives rise to floristic diversity and, ultimately, is another source 
of biodiversity. The soils at the two ends of the GLGS range are quite different. The great range of 
host rocks, elevation, latitude, and monsoon conditions within the GLGS gives rise to considerable 
soil diversity, hence contributing to the region's biodiversity. 

Hypsography and Landform Analysis 

The size of physical geographic structures in the Hengduan Mountains is large and the com- 
ponent ranges or ridges can extend hundreds of kilometers. The GLGS are the most southerly 
reaching of the major ranges. Each range can have many names where it crosses ethnic boundaries. 
These names will be given from north to south and will be abbreviated to that shown in brackets. 
(1) the most easterly transverse ridge being the Ninjingshan-Yunling-Qingshuilangshan (Yunling 
Mountains), which form the eastern bank of the Lancang River; (2) between the Lancang and the 
Nujiang Rivers is the Taniantawenshan-Nushan Ridge (Nushan Mountains); and (3) the most west- 
erly ridges of the Hengduan Mountains are the GLGS, with the "NE-SW Trending Ridges" of 
Jiangao Mountain extending into the Kachin State of Myanmar. The Shanngwa Range west of the 
Nmai and Tamai Nmai Rivers is not considered part of the Hengduan Mountains. Neither is the 
next ridge beyond the Mali River, the Kumon Range. The transverse ridge joining these Myanmar 
ranges in the north (the Patkai Mun Range between upper Myanmar and Assam) is not part of the 
GLGS. 

Northern Limit of Gaoligong Shan. — The upper GLGS are separated from the eastern 
Himalayas by a major tributary of the Yarlung Zangbo River (Brahmaputra), the San Qu River 
(Luhit River in Assam, also known as the Sang Qu and Zayii Qu River further north). This tribu- 
tary extends as far as the Zayii County and Zogang County border. A few kilometers to the north 
of the Zayii Qu rise the Parlung River and Yi'ong Zangbo River, other tributaries of the Yarlung 
Zangbo River (Brahmaputra). North of this area, the Nushan Mountains loop over the northern end 
of the GLGS. The most southerly of the mountain ranges of the Tibetan Plateau, the 
Nyainqetanglashan, are just north of the Yarlung Zangbo River (Brahmaputra). These are separat- 
ed from the GLGS by a river that is a tributary of the Nujiang River to the east. This tributary 
almost joins the drainage of the Parlung and Yi'ong Zangbo Rivers south of Bomi Town to the 
upper course of the Sang Qu River (Luhit). These mark the northwestern river boundary of the 
GLGS. Together they form a high valley between the Nyainqetanglashan and the GLGS, near 
Baxoi Town, well south of Qamdo. At Baxoi Town, the most southerly road from Sichuan Province 
and Lijiang City, Yunnan passes into eastern Zayii County towards towns of Gyigang and Zayii. 
This is just north of the Diphu Pass above the headwaters of the Tamai Nmai River. The Diphu Pass 
marks the boundary of the Patkai Range and the GLGS. All these structures combine to make a 
break in the GLGS that clearly separate it from its higher neighbors. Many of the peaks in the upper 



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Figure 1 1 . Perspective view of the Gaoligong Shan from the north look- 
ing towards the southeast. Main ridgehne shown as a yellow line. 



Nushan Mountains and eastern 
Himalaya are 7000 m plus. These 
physical features can be seen in 
the elevation perspective model 
(see Figs. 11-13 and the satellite 
image, Fig. 14). 

Southern Limit of Gaol- 
igong Shan. — The southern 
boundary of the GLGS is clearly 
defined. The GLGS end where 
the main ridge ends in an encir- 
clement by the Nujiang River to 
the east, south and west, and its 
tributary the Supa River. The 
Supa River rises off the north- 
west side of the main GLGS 
ridge on Daxue Mountain- near 
the village of Zhen'an. The Supa 
River runs northeast of the city of 
Longling to join the Nujiang 
River north of Pingda. This tribu- 
tary is almost met by the 
Mangshi River, a tributary of the 
Longchuan and Irrawadi Rivers, 
which rises just southwest of 
Longling Town. To the west of 
this promontory are the NNE- 
SSW ridges of the low mountains 
within Luxi County that meet the 
GLGS ridge just north of 
Longling Town. These are sepa- 
rated from the GLGS by a saddle 
between the aforementioned 
rivers. These physical features 
can be seen in the elevation per- 
spective model (Figs. 15-17) and 
satelhte image (Fig. 18). 

Land Form of Gaoligong 
Shan. — The elevational nature of physical features is the result of the interplay between tectonic, 
geologic, and natural erosional forces. High mountains are formed by tectonics and are maintained 
either by hard rocks and slow erosion, ongoing tectonic actions, or both. Harder rocks make for 
steeper slopes and more bare outcrops. Rain, wind, and ice work together with gravity to reverse 
tectonic uplift. The steep mountains in this area result from ongoing uplift, and the hard nature of 
their rock. Despite the fact that rainfall in the area is considerable and biotic productivity high, soil 
formation and rock decomposition are unable to wear down the mountains fast enough. 

As noted above, the majority of high ground is formed from the Precambrian rocks, mostly 
metamorphics. The main ridge of the GLGS runs due N-S and is composed primarily of these 




Figure 12. Close-up perspective view of the northernmost Gaoligong 
Shan (GLGS) looking from the west looking northeast, note the valley of the 
Luhit-Sang Qu incising into the GLGS and forming a near connection of the 
Luhit-Sang Qu-Parlung Zangbo and Nujiang drainages. Main ridgeline shown 
as a yellow line. 



CHAPLIN: GEOGRAPHY OF THE GAOLIGONG SHAN, SOUTHWEST CHINA 



541 



rocks. To the east of the ridge is 
the Gaoligong Shan Fault, which 
forms the bed of the Nujiang 
River (see Fig. 4). The action of 
the fault causes enough breccia- 
tion and mylenation of the coun- 
tr\ rock to enable the river to 
carve a gorge that is thousands of 
meters deep and form a non-gla- 
cial "U-Shaped" valley. The east- 
erly side of the Nujiang River is 
the Qintang Terrane composed of 
primarily younger rock. It also 
has a large fault structure river 
complex along the course of the 
Lancang River (see Figs. 3^). 

The non-glacial "U-Shaped" 
valley of the Nujiang River is 

formed by uplift. The land is currently being uplifted more in the west. This forces the river towards 
its east bank and hence undercuts it, thus, widening the valley floor. On the western bank in many 
places can be seen a bench that forms about halfway down from the ridge (personal observation, 
but it also can be seen on the elevation models; this bench is just visible in Fig. 15.). Most of the 
roads and towns of the Nujiang River valley are located on this bench and on the western bank in 
general. 




Figure 13. Close-up perspective view of the northemmosi Gaoligong 
Shan, showing the area included in the GLGS in green. Main ridgeline shown 
as a yellow line. 




Figure 14. Space Imagery, taicen looking ENE along the Luhit-Zayii and ScUil: (Ju Rivers towards the Nujiang. Note the 
treeline at the headwaters of the Dulong in the south fright). Image courtesy of the Image Analysis Laboratory, NASA 
Johnson Space Center (NASA 2004c). 



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Figure 15. Perspective view from tlie southeast looking along the drainage 
of the Nujiang. Main ridgeline shown as a yellow line. 



Shape of the Main 
Gaoligong Shan Ridge. — The 

mountains of the GLGS and 
southern Hengduan Mountains in 
general rise from the south to 
reach impressive heights in the 
north. Just north of the GLGS 
across the Luhit-Zayii Qu-Sang- 
Qu drainage, is a large glacier 
field with peaks rising above 
5000 m. This glacier is shrinking. 
The main GLGS ridge changes 
height from the 3001 m peak of 
Daxue Mountain Longling 
County at the southern terminus, 
down through a small saddle at 
1930 m the lowest point of the 
ridgeline, to rise steadily 
upwards to 4500 m proceeding 
north as shown in Figures 19 and 
20. The maximum height of any 
peak in the study area is 6318 m 
southeast of Zayii County. The 
course of the Nujiang River by 
comparison rises only 1500 m 
from 600 m in the south to 
around 2100 m towards the north 
of the study area. Therefore, the 
depth of the channel is much 
greater in the north than it is in 
the south. The valley is more 
than 3000 m deep at most points 
in the north and almost always 
more than 2000 m deep through- 
out the GLGS. 

The GLGS ridge is traversed by only a few passes. In the south is the pass to Longling County 
that is the main route to the Myanmar border. Luoshuidong Pass, between Bawan and Tengchong 
Towns, provides another vehicle route to Myanmar. Pianma Pass is near Lushui Town. Other pass- 
es include the E'ga Path just north of Lukiu Town, the Yaping Path north of Fugong Town, and the 
Dazhu Path a little south of Gongshan town. A full vehicle road has been built through the high 
Dulong Pass in the north. Lastly, the Zayii County and Zogang County border road from Sichuan 
to Lhasa forms the northern limit of the GLGS. 

Cross-sections through the Gaoligong Shan. — The cross-sectional profiles of the 
GLGS area were taken from the low point at the easternmost edge of the study area in the west to 
the Nujiang River in the east. These profiles are remarkable for showing the steep walls of the 
peaks and deeply incised river valleys. The northernmost profile (Fig. 21) from the Zayii River to 
the Nujiang River shows uniformly high ground. The cross-section of Bingzhoulou Town, 




Figure 16. Close-up perspective \ie\\ ot the southernmost Gaoligong 
Shan. GLGS main ridgeline shown as a yellow line. 



CHAPLIN: GEOGRAPHY OF THE GAOLIGONG SHAN, SOUTHWEST CHINA 



543 




J W^b>^ . A iHIIIMIll- ""' <1 
Figure 17. Close-up perspective view of the southernmost Gaoligong 
Shan, showing the study area included in the GLGS in green. 



Gongshan County (Fig. 22) has 
some interesting features. To the 
east is the land that is lower in 
the upper Nmai-Irrawadi Rivers 
and generally lower across the 
eastern extreme of the Himalaya 
Block. The harder rocks of the 
tectonically deformed Lhasa 
Terrane stand very high. The 
Dulong River cuts a swathe 
through them almost as deep as 
that of the Nujiang River "s val- 
le}" on the other side of the main 
GLGS ridge. To cut this deep, 
there may be another fault active, 
although it is not shown on 
small-scale maps available. The 
Nujiang River runs in a very 
deep and steep valley at this 
point. 

The next profile at Lukiu 
Town (Fig. 23) is across the area 
where the rocks are more uni- 
form Paleozoic and form the 
"NE-SW Trending Ridges" that 
extend towards Jiangao Moun- 
tain from the main ridgeline. The 
main GLGS ridge stands higher 
and there is little penetration of 
the rivers that here tend to run 
either north or south from the 
■'NE-SW Trending Ridges". 
There is little in the way of river 
erosion in the mountains here 
because of their smaller catch- 
ment basins. Although the 
Nujiang River flows in a steeper 
valley than at its outlet, it is at an 
elevation little changed from its 
exit from the GLGS. The most 
southerly profile, through the top 
of Tengchong (Fig. 24), is from 
where the GLGS has been 
opened by N-S trenches along 
which flow the Longchuan River 
to the east and the Dayang River 
in the West. The southern profile is longer and lower than the rest and shows a series of peaks and 




Figure 18. Space Imagery, taken looking east along the Longchuan River 
towards the Gaoligong Shan Ridge. Image courtesy of the Image Analysis 
Laboratory, NASA Johnson Space Center (NASA 2004b). 



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valleys where the tributaries of the Irrawadi 
River have cut into the Tengchong County 
region. These rivers also arise within the GLGS 
so have less volume or down-cutting potential. 
Uplift in this area is mostly lower than it is fur- 
ther north or it has been reduced by extension. 
The Nujiang River in the south is at about the 
same elevation as the Burmese Plain. 

Slope and Slope Direction or 
Aspect. — What is unusual about the GLGS 
area profiles is that the deep valleys are cut into 
rocks able to support steep slopes. The average 
slope angle for the whole study area is high and 
is much higher in the north (see Fig. 25). There 
are very few flat areas of large size in the 
GLGS. 

The N-S trending mountains of the main 
ridge of the GLGS together with the "NE-SW 
Trending Ridges" in the lower part both have 
unusual face aspects (see Table 1). Aspect is 
determined from the average direction that a 
slope faces relative to the sun. Throughout the 
study area, there is a paucity of north or south- 
facing slopes. The east-facing slopes are small- 
er than those to the west because they are steep- 
er. There are many facing to the west and north- 
west and then again to the northeast and then 
east. 

This fact, combined with the angle of the 
slope, means that many of the surfaces in the 
GLGS receive lower intensities of insolation 
relative to an ideal "suntrap." Maximum inso- 
lation is received on a slope that is facing south 
and that is raised to the same azimuth as the 
sun; in the GLGS this is about 25 degrees. The 
low energy capture seen in the GLGS is 
because not much of the energy of the sun is 
trapped by the slopes that face away from the 
sun. When the sun shines on a surface that is 
steeply inclined and angled away from the sun, 
its energy is dissipated over a much larger area. 
The steep terrain will also affect the local sun- 
rise, or sunset, or both, especially in a "U- 
shaped" valley. Therefore, the day length of 
direct sun. and hence the biotic productivity. 
will be much curtailed in the steep, "U-Shaped" 



See Fig 




See Fi2. 21 



r^ 



Figure 19. Plan of cross sections. 
in2 figures 21ou2h 24 



See also accompany- 



valley bottoms. These factors produce temperate 



conditions, which prevail further south in the GLGS than in most places in the world. The magni- 



CHAPLIN: GEOGRAPHY OF THE GAOLIGONG SHAN, SOUTHWEST CHINA 



545 



Ridge Profile 



'Nm*'^^ 



Zayu Profile 



3X0 



6000 



5000- 



4000- 



3000 



2000 



ICOCOJ 2CCO:0 30(XCO VXXXX) sooco 6acoo 




Meters 



Bingzhongluo 



20000 40000 60000 

meters 
Liuku Profile 



80000 




50000 



100000 



150000 




meters 
Tengchong Profile 




meters 

Figure 20. Profile of the main Ridgeline. 
Figure 21. Cross section at Zayii. 
Figure 22. Cross section at Bingzhongluo. 
Figure 23. Cross section at Liuku. 
Figure 24. Cross section at Tengchong. 



50000 



100000 



150000 



meters 



tude of this effect can be calculated using advanced GIS analy- 
sis but this is too detailed to be carried out here. 

Landforms and Biodiversity. — The unusual physical 
features of the GLGS and their great latitudinal and elevation- 
al range provide for the easy maintenance of biodiversity. The 
north-south conduit enables exchange with the high mountains 
and plateau to the north. This provides a corridor for temperate 
animals to migrate southward during harsher conditions. The 
high elevation equally acts as a barrier for warm-adapted 
organisms seeking to migrate from west to east or vice versa. 
The deep, large rivers also act as barriers. The lack of south- 
facing slopes and the deep valleys combine to make the area 
relatively more temperate than land at similar latitudes. Cold 



Table 1. Aspect faces of the GLCS. 



Aspect Direction 


Plan Area % 


North 


5.5 


Northeast 


13.0 


East 


15.5 


Southeast 


12.7 


South 


11.2 


Southwest 


13.5 


West 


16.8 


Northwest 


11.8 



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Legend 

Flat(-1) 
North (0-22.5) 












WV^S; 




Northeast (22 5-67.5) -^fs'p- 
East (67.5-112.5) 
Southeast (112.5-157.5) 
South (157.5-202.5) 
Southwest (202.5-247.5) 
West(247.5-292.5) 
Northwest (292.5-337.5) 
North (337.5-360) 



Figure 25. The plan \iew showing the A\erage Slope of 
the Gaolisono Shan. 



Figure 26. The plan view showing the Aspect of the 
Gaohgong Shan 



air can flow into the valleys from high ground surrounding them. These "frost traps" lead to fre- 
quent fogs and temperature inversions. The high hills with damp air coming from the west have 
significant amounts of rainfall on their western slope. These conditions can lead to Foehn heating 
as damp air is forced over the ridge by the prevailing southwesterly winds. The unusual physical 
features combine to multiply the number of opportunities for microclimates. Furthermore, these 
physical features are not fixed in time but are dynamic due to the nature of the underlying geolog- 
ical processes. This dynamism provides ample opportunity for adaptation and vicariant events to 
further promote biological diversification. 



CHAPLIN: GEOGRAPHY OF THE GAOLIGONG SHAN. SOUTHWEST CHINA 



547 




Figure 27. Drainage pattern of major rivers through the Hengduan 
Mountains emphasizing the interdigitated nature of the drainages. Data from 
HydrolK(USGS2000). 



Hydrology 

The large N-S flowing rivers 
of the Hengduan Mountains are 
of major importance to East and 
Southeast Asia. The rivers are 
long, stretching from the Tibetan 
Plateau to three different seas, 
the Yellow Sea, the South China 
Sea. and the Andaman Sea in the 
Bay of Bengal. Because of their 
length, each one of the rivers can 
have many names where they 
cross ethnic boundaries. These 
names will be given from north 
to south and in this paper they 
will be abbreviated to those 
shown in brackets, which are 
their names as used in the GLGS 
region. These are not necessarily 
the rivers' more widely used 
common English names. When 
using the abbreviated name the 
intention is for the reader to think 
of the whole drainage not just 
that portion in western Yunnan. 

From east to west the main 
rivers are the Wulanmulunhe- 
Muluwusuhe-Tongtianhe-Jinsha- 
Cang Jiang-Yangtse River 
(Jinsha River), the Lancang- 
Mekong River (Lancang River) 
and the dNgul-chu-Naquehe- 
Nujiang-Thanl win- Sal ween 
River (Nujiang River). An impor- 
tant tributary of the Nujiang 
River is the Nanding River, flow- 
ing just below the southern end of the GLGS. Starting in China and flowing into Myanmar are the 
south-southwesterly flowing tributaries of the Irrawadi. They are, from north to south, the Dulong- 
Taron River joins the Nmai Hka River (Dulong River), Dayang River, Wanding River and 
Longchuan-Shweli Rivers (Longchuan River). In the southern part of western Yunnan, just to the 
west of the GLGS, is the source of the Lishehe-Yuanjiang-Hong River (Red River) rising between 
the Lancang and Jinsha Rivers and its large tributary is the Black River. In the north are the Yarlung 
Zangbo-Brahmaputra River (Yarlung Zangbo River) and its two easternmost tributaries the Luhit- 
Zayli Qu-Sang Qu River (Sang Qu River) and the Yi'ong Zangbo River and Parlung Zangbo River. 
These form the northwest border of the GLGS. They flow to the Bay of Bengal in the west. 

It must be strongly stressed that the drainage pattern around the Hengduan Mountains is com- 




FiGURE 28. The modern drainage catchment basins data from Hydro IK 
(USGS 2000). 



548 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 28 

plexly interleaved. This complicated pattern interdigitates to cause formidable barriers (see Fig. 
27). To the north the Yarlung Zangbo River (Brahmaputra)and its easternmost tributaries form a 
natural barrier. The Yarlung Zangbo River (Brahmaputra) loops in a big bend from flowing east for 
hundreds of kilometers to turn south, then west, and on to flow southwest to the sea. This loop 
forms a major obstacle to migration or dispersal. Any migration from the west would be caught in 
this big bend and would have to back track or go north to get around it. From about the 10 Mya, 
the climate and ecology to the north would have been very different than that in the big bend area 
(Jablonski 1998). Today the barrier is complete as the ecology changes from humid tropical to tem- 
perate to alpine tundra within a few kilometers along the hills of Arunchal Pradesh. The Jinsha 
River makes a similar big bend in the opposite direction flowing to the south then east, and then 
north to turn eventually east again to the Pacific. Between these two, the Yarlung Zangbo 
(Brahmaputra) and Jinsha Rivers, the following Rivers, Lancang, Nujiang, the Dulong Nmai, and 
Irrawadi proper, all flow north-south. To the east, starting near the big bend of the Jinsha River is 
the source of the Red River and to the west of the Red River is its tributary the Black River that 
runs parallel to it. The Black River rises near to the Lancang River just east of the GLGS. Further 
east of the source of the Red River is the source of the Pearl River. These form NW-SE river 
drainages that cut off the approach to the Hengduan Mountains from southern China. The approach 
from due south to the GLGS (but not the Nushan Mountains) is cut off by the Lancang River and 
Nanding River and by the eastward flowing section of the Nujiang River. The approach to the 
Hengduan Mountains from the southwest is blocked by the Nujiang River and the Shweli- 
Longchuan River, Dayang River and other tributaries of the Irrawadi River. The approach from the 
west is blocked by the Yarlung Zangbo River (Brahmaputra), Irrawadi River, Tamai Nmai River, 
and the Dulong Nmai Rivers. West to east migration would be the most difficult because of the 
need to cross rivers and change elevation across the ridges of western Myanmar. The most isolat- 
ed of the Hengduan Mountains ranges is the GLGS. The GLGS is highly isolated by its almost con- 
tiguous surrounding rivers. 

The rivers of the Hengduan Mountains make them exceedingly good biological barriers to ter- 
restrial organisms (Mackinnon et al. 1996). For many terrestrial organisms, migration into the 
GLGS or dispersion from them is very difficult. Fording the rivers is not easy because they run in 
very deep, precipitous valleys cut into the mountains. The ridges of the Yunling Mountains, Nushan 
Mountains, and the GLGS are steep, and traversing them requires agility and considerable environ- 
mental adaptability. The steepness of the riverbeds makes the current strong; some of the rivers 
have dangerous category five rapids. The rivers also carry high volumes of water and experience 
occasional catastrophic floods. 

Biodiversity Implications of the Hydrology Network. — It can be seen from Figure 28 
that within the GLGS very little of the area belongs to the Nujiang River watershed. Most of the 
land area falls within the IiTawadi system. This can be inferred from the ridge profile as well. The 
Irrawadi is a much newer system than the Nujiang River because it does not drain north of the con- 
tact zone with India. A number of factors influence aquatic diversity, including age, temperature, 
and current. 

In the past, these rivers would have been even more of a barrier than are today. The evidence 
strongly suggest that the larger rivers predate the closing of the Tethys Sea. The present rivers and 
the paleo-rivers drained regions as far north as the Kunlung Mountains an area north of the paleo- 
shore of the Tethys Sea.. Before the formation of mountains, there was no rain shadow. The moun- 
tains rose in sequential thrust belts developing in the west of the GLGS as the Indian Plate impact- 
ed the Eurasian Plate. Therefore, the paleo-rivers would have had to drain the area that is now 
behind the Himalaya. The area to the north is now in the Himalayan rain shadow. The rain shadow 



CHAPLIN: GEOGRAPHY OF THE GAOLIGONG SHAN, SOUTHWEST CHINA 549 



was absent throughout most of the history of these rivers. Therefore, they would have captured 
larger volumes of water than the impressive amounts they do today (see Fig. 28). The final uplift 
of the Himalaya to their current elevations has been in the last 7-3 million years of the 55 million 
years since India first contacted Eurasia. It was not until after this time that the rain shadow was 
extensi\'e enough to cause aeolian erosion and loess started to be blown from the rising Tibetan 
Plateau. The main north-south flowing rivers divide the area biogeographically and socially. The 
antiquity of the rivers has insured that the areas divided by them accumulated considerable pre- 
Neogene diversity. Their large size and long length provide opportunity for aquatic diversity to 
evohe in multiple habitats. The changing levels in the rivers gave rise to fast currents and rough 
water, which have limited aquatic diversity and prevented migration and dispersal of endemics, 
while opening the possibility of local adaptation. Their encirclement of the GLGS has created one 
of the most isolated regions in the world, with a high number of endemic species. Another, biolog- 
ically significant aspect of the large rivers is that they have provided unusually deep and secluded 
valleys, which have acted as refugia. Species can move up and down elevational gradients to main- 
tain thermal equilibrium during periods of rapid temperature fluctuation. The unusual climates of 
the river valleys have promoted the successful survival of species extirpated elsewhere. The depth 
of the valleys effectively limits biotic productivity through reduced insolation relative to latitude 
and high humidity reduces light levels further. The rivers buffer extremes of temperature due to 
cloud and fog formation from high humidity. The river valleys allow warmer wind from the south 
to penetrate far to the north during the winter monsoon. 

Definition of Gaoligong Shan 

Previous Definitions and Biogeography 

The GLGS have been previously defined by several workers and environmental organization. 
Some definitions use the physical features to define the area. The best previous definition of the 
GLGS is that of Li: "The Gaoligong Shan is: the mountain range between Nujiang River and 
Irrawadi River, it is located in N 24°40'-28°30', covering totally 111,000 square kilometers, which 
includes the whole territory of Tengchong County, part but most of Longling, Baoshan. Lushui, 
Gongshan County area, besides N Burma area (Kachin State)" (Li 2000:vii). Although, this defini- 
tion has a straight-line, latitudinal cutoff in the north and omits the SW ridge in Yingjiang County 
it covers most of the GLGS as it is defined in this paper. Lan and Dunbar define the Gaohgong 
Shan Region differently: "The region referred to as Gaoligong Shan here includes all lands west of 
the Salween (Nujiang) River in Yunnan. The entire region is situated at the southern edge of the 
eastern Himalayas, the westernmost region of Yunnan Province, and in the western part of the 
Trans-Himalayan Mountains" (Lan and Dunbar 2000:275-276). In practice, however, they used an 
essentially political definition; therefore, all of the land in Myanmar including the interconnected 
"NE-SW Trending Ridges" and the territory on the western side of the main GLGS main ridgeline 
in Lushui and Fugong Counties were excluded. Although politically this was not unreasonable, nat- 
ural phenomena do not follow political constructions. The World Heritage listing definition of the 
Three Parallel River Region of Yunnan includes only the northern part of the GLGS ending at 
Lushui County at the end of the very highest ridgehne. It emphasizes the role of the gorges more 
than that of the mountains (UNESCO 2003). The GLGS are included in the recently revised and 
corrected definition of "Mountains of Southwest China, Biodiversity Hotspot" used by 
Conservation International (Conservation International 2005). However, it includes the whole 
Hengduan Mountains and Longmen Mountains and so is of little use in discussing just the GLGS. 



550 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

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Other workers defined the area according to biogeographical or ecological considerations. The 
Hengduan Mountains subalpine conifer forests zone (PA0509) used by the World Wide Fund for 
Nature (WWF) does not extend so far south or west as does the GLGS (Carpenter 2001a). The part 
of the GLGS region is included in Nujiang River Lancang Gorge Alpine Conifer and Mixed Forests 
(PA05 16). "The Nujiang River Lancang Gorge ecoregion includes the valley system through which 
rivers flow down from the Tibetan Plateau into the tropical hills of northern Indochina" (Carpenter 
2001b). This definition missed the western slopes of Myanmar that are in Northern Triangle sub- 
tropical forests (IM0140) (Than et al. 2001). The GLGS are spread between three different eco- 
zones according to the WWF. Similarly, the GLGS are split by many other biogeographers and 
ecologists. Because of its elevation and latitude, the northern part of the GLGS is often classed as 
part of the Tibetan Plateau; the middle reaches with Yunnan Plateau; and the southern parts as sub- 
tropical forest continuous with that of Myanmar or Thailand (Mackinnon et al. 1996; Zhao 1986). 
The area of western Gongshan County and Fugong County is included in the Himalayan Southern 
Slope Region by Zhao (1986). Mackinnon (1996) includes most of the GLGS in the Paleartic 
Realm, Southwest China Province but makes a new subunit for the Nujiang River Lancang Gorges 
Area 39f, the middle of the GLGS is within sub-unit Yunnan Plateau 39a, whereas, the south is in 
the Indo-Malayan Realm, Tropical South China Province, sub-unit 10 the Thailand Subtropical 
Monsoon Forest (Mackinnon et al. 1996). 

The Definition of the GLGS as Used in this Paper 

The name GLGS refers to mountain features, so it is best that it is defined by its physical geog- 
raphy. Therefore, the GLGS comprise the contiguous mountain ridges between the drainages of the 
Nujiang River (Salween River) and the Irrawadi River systems. In the north beyond the Irrawadi 
River headwaters the GLGS are between the Sang Qu River (Luhit), a tributary of the Yarlung 
Zangbo River (Brahmaputra) and the Nujiang River (Figs. 12-14). 

The contiguous ridges were defined as land over 1 800 m. The areas above 1 800 m form inter- 
connected ridges that join the Hengduan Mountains. This elevation was chosen as it is the cutoff 
of the "Monsoon Evergreen Broad-Leaves Forest" belonging to the Castanopsis hystrix and 
Castanopsis echidoncarpa forest type. This forest is distributed in moist ravines, on the east-fac- 
ing slope in the southern part of the region, at elevations rising to but not above 1800 m (Li 2000). 
Using "Monsoon Evergreen Broad-Leaves Forest" for choosing the elevation for the ridges was 
helpful for two reasons. First, there are no barriers between "Monsoon Evergreen Broad-Leaves 
Forest" within the Gaoligong Shan and the same zone that spreads throughout a large area to the 
south and east covering broadly most of Myanmar and much of Southeast Asia. Second, above 
1800 m the ridges are complete and continuous within the GLGS (Fig. 29). 

In areas where there is neither a river barrier nor an extending ridge, the study area was cur- 
tailed at the 1000 m mark. This was necessary for only two small areas to the southwest of the "NE- 
SW Trending Ridges." Here the essentially flat area north of the Dayang River and south of the 
next tributary of the Irrawadi extend far into Myanmar before joining the Irrawadi. This area is also 
extensively farmed. 

The Dayang River was followed as the boundary in the south until it turned to the north near 
Nansong Town. Then the boundary was cut across the top of Lianghe County from Nansong to 
Pingshan and the Longchuan River. The Longchuan River provides the boundary of the GLGS 
until it reaches the western slope of the GLGS ridge. Here the Longchuan River turns north along 
the border of Tengchong and Longling Counties. The final section of the GLGS southern border is 
encompassed by a line following the lowest contours round the end of the Mangshi River until it 



CHAPLIN: GEOGRAPHY OF THE GAOLIGONG SHAN, SOUTHWEST CHINA 



55. 



reaches the first tributary of the Nujiang River 
the Supa River. The Supa River rises north of 
Zhen'an Village and runs from the northwest 
slope of Daxue Mountain on the GLGS ridge to 
travel west, before traveling south, southeast, 
and then eventually east to encircle the south- 
em point of the GLGS main ridge. This defini- 
tion provides the shortest route between the 
Nujiang River and Irrawadi drainages that 
encompasses the entire ridge complex. 

As per the discussion of the GLGS above, 
the low ridges to the southwest of the GLGS, 
low ridges in Yingjiang Lianghe Counties and 
those in western Longling that extend into Luxi 
Count}' could be argued to be also a part of the 
Gaoligong Shan. However, the decision not to 
include them was based on hypsographic argu- 
ments alone. The ridges are not contiguous but 
are separated by low points or watershed 
boundaries. These lower ridges all belong to 
the Irrawadi system not the Nujiang River 
drainage system. Therefore, they lie outside of 
the watershed between the Nujiang River and 
Irrawadi. The Nujaing-Irrawadi drainage 
boundar}' is at the Longchuan River, which 
forms the border between Longchuan, Lianghe, 
and Luxi Counties and this boundary is north of 
those County's southernmost ranges. 

The Nujiang River and Irrawadi River 
Valleys provide the cutoff points to the 
Gaoligong Shan as these are the lowest points. 
The GLGS are defined as a hypsographic fea- 
ture. On the opposite bank of these river 

drainages the slope must, by definition, rise again. Therefore, the contiguous slope runs only 
between the rivers. 

The main observations about the physical features of the GLGS are presented in Table 2. 




Figure 29. Map indicating all contiguous land over 1800 
m in brown. 



Conclusions and Implications for Biodiversity 



The Hengduan Mountains are a haven of biodiversity. The GLGS are the most isolated of the 
ranges of the Hengduan Mountains, due in large part to the drainage pattern. The potential of the 
GLGS for preserving biodiversity, as well as causing it, may be unique in Eurasia. The position of 
the GLGS in Eurasia enables them to be a reservoir of biodiversity for all of East Asia. 

The GLGS straddle the Indo-Malayan and the Paleartic biogeographic realms and have been 
split into different biogeographic provinces by different workers. The descriptions of these 
provinces have not caught up with modem understanding of tectonics, leading to considerable con- 
fusion. The physical geography of the GLGS is the result of tectonic activity. All of the plates form- 



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Table 2. General Facts about the Gaoligong Shan 



1. Maximum Linear Length along Main Ridge 585 km. 

2. Minimum Linear Length along NE-SW Ridge 565 km. 

3. Maximum Width 150 km in the south near Tengchong. 

4. Maximum Width 100 km in the north near Gongshan Town. 

5. Minimum Width 48 km near Fugong Town. 

6. Bounding Box 91 ATE, 29.5 1°N and 99.03°E, 24.37°N decimal degrees. 

7. Maximum Elevation 6318 m southeast of Zayii County. 

8. Minimum Elevation 183 m Drainage of the Nmai River in Myanmar. 

9. Minimum Elevation 620 m Drainage of the Nujiang River. 

10. Mean Elevation 2638 m. 

11. 62% of the land lies between 1500-3500 m. 

12. 11% of the land is above the approximate tree-line of 4500 m. 

13. Only 7.8% of the surface area is essentially flat (slope < 3%). 

14. Mean slope for the whole area including the drainages is 13.4%. 



Elevation Banc! 


Surface Area 


3d Sutface Area 


Area in Band 3d 


%ofTotal3dArea 




In Plan View knr- 


Along Slope knr- 


Surface hw- 


in Band 


0^99 


41937 


AAUl 


279 


0.63 


500-999 


41661 


43867 


1642 


3.72 


1000-1499 


40056 


42226 


4078 


9.24 


1500-1999 


36114 


38147 


8395 


19.02 


2000-2499 


28007 


29753 


8453 


19.15 


2500-2999 


19951 


21300 


6040 


13.68 


3000-3499 


14309 


15260 


4520 


10.24 


3500-3999 


10133 


10740 


3491 


7.91 


4000^499 


6891 


7249 


3334 


7.55 


4500-4999 


3763 


3915 


3112 


7.05 


5000-5499 


778 


803 


798 


1.81 


5500-5999 


4 


5 


5 


0.01 


Total Area 


41937 km-2 


44147 km-2 







ing the GLGS are from Gondwanaland. but some of them have been in contact with the Eurasian 
Plate (Paleartic Biogeographic Realm) for upwards of 200 million years. The Indo-Malayan region 
is physically an assembled unit composed of units of vastly different ages. For a review of current 
usage of China's zoogeographical zonation refer to Mackinnon et al. (1996). 

Tectonic forces themselves create a genetic "melting pot"' for biodiversity. The paleo-separa- 
tion and subsequent re-aggregation of plates from Gondwana and the collision of plates from dif- 
ferent paleo-continents laid a foundation of high genetic diversity. This was accentuated by the 
region's long and complex uplift history. Slowly rising landmasses provided opportunities for 
adaptive changes in resident organisms. Geographic isolation due to the tectonically driven inci- 
sion of the landmass by massive rivers has further enhanced biodiversity through vicariance. 

Tectonic forces give rise to a diversity of host rock and soil types. This diversity has further 
enhanced the potential for increased biodiversity. The great range of host rocks, elevations lati- 



CHAPLIN: GEOGRAPHY OF THE GAOLIGONG SHAN, SOUTHWEST CHINA 553 



tildes, and monsoon conditions within the GLGS has given rise to considerable soil diversity. This, 
in turn, gives rise to floristic diversity and heightened biodiversity at all higher trophic levels. 
Adaptive forces related to tectonics operate at both the macro as well as the micro landscape scale. 
The many preserved volcanic cones in the area of Tengchong resulting from tectonic melting, could 
be a source of local adaptation and vicariant speciation of smaller organisms as is known from 
other volcanic fields, e.g.. DwsophUa on Hawaii. 

Summary 

i^iiEis^HM^-^o mM.}A±M^m±'Amj&^K^ (JablonskiandPan 

1988) o JAm^MiJMyKW-M^'^ii (Mackinnonetal. 1996) o mkmJWm^ff^^ 

sfBjo ijmimu-^Hm^it. ^mmE.mmm^j^, wi^f^±n'M^m^^io m 
m^ ill ummm^t^^m. #wfp#^-tft!^^o Mmf^ihmm^mmjEjn'±-'\^ 



554 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 28 



Acknowledgements 

Thanks for support to Environmental Systems Research Institute (ESRI) for supplying data 
and software to the California Academy of Sciences. Thanks also to Dr. Nina G. Jablonski for 
advice and help with editing and to Dr. Peter Fritsch and Dr. Bruce Bartholemew for comments on 
an earlier draft and to several anonymous reviewers whose comments led to a greatly improved 
presentation. Thanks also to Dr. Nina G Jablonski for the invitation to visit the GLGS. Dr. Lihua 
Zhao, Department of Botany, California Academy of Sciences, prepared the Chinese versions of 
the conclusions and summary statements that accompany this paper. This publication represents 
Contribution No. 39 of the Center for Biodiversity Research and Information (CBRI) and 
Contribution No. 27 of the China Natural History Project, both at the California Academy of 
Sciences. 

Cartography and data. — Care was taken to ensure that political borders were depicted rep- 
resentatively; however, they are provided only for indicative purposes and do not represent any ter- 
ritorial claim or agreements. Representation of borders will depend on the dataset used and differs 
slightly between maps. The reader should note that the borders are disputed in a number of areas 
within the Hengduan Mountains region. No opinion is expressed or implied in the cartography or 
text. Place names are given to be as informative as possible to the general reader; they do not mean 
to imply any special meaning to the names used in this paper. Place names are taken from the Map 
of the People's Republic of China (Carto. Pub. Hse., 1984) and from common usage. 

The data used in this paper came from a variety of sources. The following datasets were used 
either alone or in combination to produce the maps (ESRI 1996; F.A.O. 2005; Steinshouer et al. 
1997; USGS 1993, 2000, 2004; Wandrey and Law 1997). All maps are original maps produced 
from the data using ArcGIS© and Arc View© (ESRI 1999, 2004). 

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Copyright © 2005 by the Cahfornia Academy of Sciences 
San Francisco. California. U.S.A. 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 

Volume 56. No. 29. pp. 557-599, 65 figs., 3 tables December 30, 2005 

The Giant Pill-Millipedes of Madagascar 

Revision of the Genus Sphaeromimus, 

with a Review of the Morphological Terminology 

(Diplopoda, Sphaerotheriida, Sphaerotheriidae) 

Thomas Wesener^ and Petra Sierwald^ 

1 Department of Animal Morphology & Systematics, Ruhr University of Bochum, 

Universitdtsstrasse 150, D-44701 Bochum, Germany: -Zoology, Insects, Field Museum of Natural 

Histoi-y, 1400 South Lake Shore Drive, Chicago, IL 60605, USA; Email: sierwald@fieldmuseum.org 

The Malagasy sphaerotheriid genus Sphaeromimus DeSaussure and Zehntner, 1902 
is revised. Known heretofore from a single male specimen, the genus now contains 
three species, Sphaeromimus musicus (DeSaussure and Zehntner, 1897), Sphaero- 
mimus splendidiis sp. nov. and Sphaeromimus inexpectatus sp. nov. The female of S. 
musicus is described here for the first time. The mouthparts of giant pill millipedes 
were observed for the first time using scanning electron microscopy and species- and 
genus-level characters are illustrated. Intraspecific variation of the female stridula- 
tory organ, the 'washboard' is described. For the first time in Malagasy 
Sphaerotheriida, some ecological comments are given. Characters found in the male 
telopods and the female stridulatory organ (the washboard) indicate that characters 
employed previously for the definition of subfamilies and tribes cannot be main- 
tained and the monophyly of such groups remains questionable. 

Keywords: Sphaeromimus, Sphaerotheriida, giant pill-millipedes, Madagascar, Diplopoda. 

Based on its species-richness and high level of endemism (Myers et al. 2000), Madagascar was 
recently listed among the eight eminent biodiversity hotspots of the world. Madagascar, as the 
fourth largest island of the world, harbors a diversity of different ecosystems, resembling in this 
regard a small continent. Due to its over 150 million years of isolation from the closest continen- 
tal landmass Africa (Rabinowitz et al. 1983; Wells 2003), its flora and fauna are unique and very 
distinct from that of other regions of the world. Furthermore, the fauna and flora of Madagascar are 
extremely poorly known, as is the case for many species-rich regions outside northwestern Europe 
and North America. Ongoing faunistic research on the island of Madagascar continues to discover 
numerous new species, even among vertebrates (Jenkins 1993; Glaw and Vences 1994; Sparks and 
Stiassny 2003). Since the destruction of natural habitats is advancing on the island at an alarming 
rate, alpha-taxonomic research with regards to invertebrates is extremely urgent, as many species 
may vanish before ever being described. Three Malagasy ecosystems, the east coast littoral forest, 
highland vegetation and the western dry deciduous forests have shrunk by over 90% of their for- 
mer distributions and belong now to the most threatened ecosystems of the world (Ganzhorn et al. 
2001; de Gouvenain and Silander 2003; Vincelette et al. 2003). 

The millipede genus Sphaeromimus revised below illustrates the understudied invertebrate 
diversity as well as the threatened status of its species. The diverse arthropod class Diplopoda is 

557 



558 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 29 

one of the severely understudied animal groups. Over 9,000 species have been described so far (an 
exact species catalog does not yet exist) and estimates of millipede species richness are given as 
approximately 80,000 species worldwide (Hoffman 1980). As far as it is known, millipede species 
are often microendemic with very small distribution ranges; in some cases related species may 
occur just 20 km apart (Enghoff 1983; Hamer and Slotow 2002; Mesibov 1998) Despite the fact 
that most millipedes are macroinvertebrates (adult size from a few millimeters to 28 cm body 
length in Archispirostreptus gigas (Peters 1855)) and are of considerable ecological importance for 
litter breakdown within the decomposition cycle (Wolters and Ekschmitt 1997; Curry 1994; 
Crawford 1992; Schaefer 1990), biological research on the class suffers from lack of alpha-taxo- 
nomic attention, mainly due to the paucity of taxonomic experts for the group. 

The Malagasy giant pill-millipede genus Sphaeromimus was heretofore known from a single 
male specimen of Sphaeromimus musicus. Among recently collected material by the senior author 
and from survey work by Goodman (Field Museum) and Griswold (California Academy of 
Sciences), female specimens of S. musicus and material of two new species of the genus were dis- 
covered. The four-jointed anterior telopods are the distinguishing feature of the genus. In 
Zoosphaerium Pocock, 1895, the other genus of giant pill-millipedes occurring in Madagascar, the 
anterior telopods have only three joints. Furthermore, features of the female vulva in 
Sphaeromimus do not agree with characters used by Jeekel (1974) in the most recent classification 
of the millipede order Sphaerotheriida and further phylogenetic analyses of the order will be 
required to clarify its internal classification. 

The two newly discovered species of Sphaeromimus described below are each known only 
from small, isolated remnants of the southern littoral forest on Madagascar. This very limited dis- 
tribution, the still ongoing anthropogenic influence in these remaining littoral forest patches and 
further possible disturbance of the habitat by mining projects may make these two new species like- 
ly to be among the most endangered millipede species of the world. 

Material and methods 

The senior author (T.W.) collected specimens of the two new species described here during 
fieldwork in Madagascar in March and April 2003. Specimens of S. musicus were borrowed from 
the California Academy of Sciences (CAS) and the Field Museum (FMNH). 

Specimens were euthanized using ethyl acetate, straightened and preserved in 70% ethanol. All 
measurements are in mm. 

Dissections, illustrations. — Dissections were made with a scalpel, in very small speci- 
mens with a dissecting pin. The following structures were dissected (a) the anterior and posterior 
pair of telopods, which were separated from each other using a needle; (b) the left leg of the 9* pair 
in males and females; (c) the 2"^ leg pair in females; (d) the L' leg pair with l^t sternite in females 
and males, (e) the subanal plate with 'washboard" in females; and (f) a section of the endotergum 
from a tergite in the center of the body, removed using scissors. Dissected specimen parts were 
cleared in clove oil. Drawings were done using a camera lucida mounted on a dissecting or com- 
pound microscope depending on size of specimen. Small specimens were held in position using 
clean sand at the bottom of dissecting dishes. 

For scanning electron microscope examinations the following parts were dissected: (a) The 
right/left antennae were cut off with a scalpel near the insertion in the head, (b) The gnathochilar- 
ium was removed by cutting along its base with a scalpel and separating the tentorium with scis- 
sors. After removal of the gnathochilarium. (c) the mandibles were cut easily at the first joint with 
scissors and scalpel, (d) The epipharynx was separated from the head with a needle and then pulled 



WESENER AND SIERWALD: REVISION OF THE GENUS SPHAEROMIMUS 559 



out with forceps, (e) The remaining head capsule was separated from the body using forceps, (f) 
The 2"*^ leg coxa of males was removed from the body. 

SEM PREPAR-\TiONS. — Specimens were dehydrated through a series of alcohol to 100% 
ethanol. mounted on stubs using sticky tabs and air-dried overnight. The 2"^ leg coxa of the male 
with the gonopore was critical point-dried. Stubs were sputter-coated with gold and observed with 
an AMRAY 1810 SEM (Field Museum). 

Terms 

As is true for many millipede groups, systematic treatments of the order Sphaerotheriida are 
scant and were done by a few authors, e.g., Verhoeff (1927, 1928, German) and Attems (1897, 
German). Silvestri (1917. Latin), Jeekel (1974. English) and recently by VandenSpiegel et al. 
(2003. English). Authors used terms in the various languages and the equivalency of such terms in 
the different treatments is sometimes difficult to determine. Since millipede morphology is less 
well known than that of other arthropod groups, which in the past have been explored more exten- 
sively with high quality light microscopy and scanning electron microscopy, the nomenclature of 
several morphological terms is currently neither standardized nor stabilized in the Diplopoda. For 
that reason we list terms used in this paper, along with terms used by other authors for apparently 
the same structure. Our use of such terms does not necessarily imply homology. 

Anal shield. — Formed by the fused tergites of the last 3(?) diplosegments (= pygidium of 
authors, e.g., VandenSpiegel et al. 2003). In males of some sphaerotheriid species the anal shield 
is invaginated in the middle (Fig. IIH). Such invagintion may play a role in mating behavior. 

Antennae. — The first visible joint of the antennae, inserting in the antenna socket, is termed 
1*' antennomere. 

Anterior paratergite depressions. — Denotes the anterior rim of the lateral extensions of the 
tergites (Paratergite, see below), a well circumscribed slightly concave area which glides under the 
posterior margin of the proximal tergite during volvation (Figs 1, 28-29, 50). Recent treatments on 
Sphaerotheriida did not explicitly discuss this morphologically distinct area. 

Bursa. — Jeekel (1974) applied this term for the structures of the female vulva below the oper- 
culum. The 'bursa' consists of two sclerites, the exterior and inner plate (EP and IP) (Figs 5, 33, 
55j. 

Endotergum. — The underside of the posterior margin of the tergites carries crenulations, 
spines and bristles, often in a species-specific arrangement (see VandenSpiegel et al. 2003; 
•Unterblatt' sensu Verhoeff 1928: plate 10, fig. 123). 

Gnathochilarium. — Since homologies with sclerites of helminthomorph gnathochilaria are 
unresolved (see Hoffman 1976:125), the sclerite terminology used here for sphaerotheriid 
gnathochilaria is descriptive. Usage of the term lamellae linguales below does not constitute a 
statement of homology. 

Harp. — A set of ridges located on a discrete plate on the first joint of the anterior telopods of 
males (Figs 8, 37, 58). 

Imier horns of posterior telopods. — Lobe-like projections attached mesally to the syncoxite. 
termed coxal horn by VandenSpiegel (2002), and 'Homer des Syncoxit' by Verhoeff (1928:676). 
Indicated here by IH = inner horn. Also, see below under 'Telopods' (Figs 11, 35, 57). 

Lamellae linguales. — Two longitudinal sclerites between the left and right paramentum of the 
gnathochilarium. In Sphaeromimus, the two sclerites are partly fused. At the distal tip of the lamel- 
lae linguales are pads carrying sensorial cones (Fig. 48). These pads were termed 
'Zapfchenkappen' by Verhoeff (1928:872). The homology of these two sclerites with the lamellae 



560 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 29 

linguales of the Helminthomorpha is questionable (see Hoffman 1976). 

Male gonopore. — Opening of the vas deferentia on the posterior wall and the inside margin 
of the coxa of the 2"^ leg pair (Fig. 27). Verhoeff (1928:695) stated that in Sphaerotheriida the male 
genital opening consists of a small, inconspicuous pore; other authors (e.g., VandenSpiegel et al. 
2003) described somewhat more complex structures and denoted them with the terms penes and 
pseudopenes. DeSaussure and Zehntner (1897/1902) illustrated the male gonopore in several 
species and genera of the Sphaerotheriida showing different structural elements. 

Molar plate process. — Elongated process attached to the upper side of the molar plate towards 
the roof of the head (Figs 18, 40), as it occurs in the millipede clade Pentazoniida (comprising the 
orders Glomerida, Glomeridesmida and Sphaerotheriida). This structure is very prominent in 
sphaerotheriids and can be found on illustrations of other pentazonid mandibles (e.g., in glomerids 
by Kohler and Alberti 1990, fig. 2-3; in sphaerotheriids by Silvestri 1917, fig. 2). No term has been 
coined for this structure. 

Paratergite (Verhoeff 1928:385, German). — Lateral extensions of the tergites. The anterior 
paratergite depressions (see above) are located on the dorsal side of the anterior margin of the lat- 
eral extensions of the tergites (Figs 1, 28-29, 50). Verhoeff (1928:385) also used the term 'Seiten- 
lappen.' These lateral tergite extensions were sometimes called paranota. Paranota is commonly 
used for the metazonite extensions in Polydesmida. Since the latter denotes a different anatomical 
part than the 'Paratergite' sensu Verhoeff in the Pentazonia. we prefer to call the structure lateral 
tergite extensions or paratergite. 

Sensorial cones. — Myriapods feature a variety of sensorial structures, one of which are cones 
with a small pore on their tip. All such structures called sensorial cones in this paper have this par- 
ticular anatomy. The distribution of such cones may reveal species- or genus-specific characters. 

Subanal plate. — Hypoproct or ventral scale of authors, equipped with a stridulatory organ 
(washboard) in females of Sphaeromimus. 

Telopods. — In the Pentazonia. males have two pairs of modified legs, the anterior and poste- 
rior telopods, at the end of their bodies. These telopods are involved in mating behavior and sperm 
transfer. It is commonly assumed that these are homologous to walking legs and thus the most 
proximal joint is called the coxite. In Sphaerotheriida, the coxites of each telopod pair are fused, 
forming a 'syncoxite.' The homology of the more distal joints with podomeres is uncertain. Here, 
the joints distal to the syncoxite are indicated by numbers 1-3 (posterior telopods) or 1^ (anteri- 
or telopods) respectively. Some authors (Mauries 2001) distinguish between the anterior and pos- 
terior telopod by using the terms 'paratelopod' (anterior telopod) and telopod (posterior telopod). 

Thoracic shield ('Brustschild' sensu Verhoeff 1928:473). — Formed by the enlarged tergite of 
the 2"'! body segment, the one following the collum. It features wide lateral lobes with a distal con- 
cave groove ("Gruben des Brustschildes' sensu Verhoeff 1928:473) and a conspicuously raised 
brim, involved in volvation (Verhoeff 1928:473). 

Vulva. — The vulva consists of the bursa and the operculum. Many authors used the term 
'cyphopods' for the female organs in millipedes. 

Washboard. — A stridulatory apparatus termed washboard by Jeekel (1999) is located on the 
subanal plate (=Hypoproct or ventral scale) at the caudal end of the body of females (Figs 7, 34, 
55). 

Abbreviations 

CAS California Academy of Sciences, San Francisco, USA 

FMNH Field Museum of Natural Histor}', Chicago, USA 

MNHN Museum National d'Histoire Naturelle, Paris, France 



WESENER AND SIERWALD: REVISION OF THE GENUS SPHAEROMIMUS 561 



12T 


12* tergite 


AI 


Anal shield invagination 


AS 


Anal shield 


EP 


Exterior plate of vulva 


IH 


Inner horns on syncoxite of posterior telopods, 


IP 


Inner plate of vulva 





Operculum of vulva 


PL 


Pleurite 


S 


Stemite 


TO 


Tomosvary organ 



Results 
Genus Sphaeromimus DeSaussure and Zehntner, 1902 

Sphaeromimus DeSaussure and Zehntner, 1902. 

Sphaeromimus, Attems 1942.— Jeekel 1971, 1974, 1999 — Enghoff 2003. 

Type species. — Sphaewpoeus musicus DeSaussure and Zehntner, 1897. Other species 
included: Sphaeromimus splendidus sp. nov., Sphaeromimus inexpectatus sp. nov. 

The Malagasy sphaerotheriid genus Sphaeromimus was first described by DeSaussure and 
Zehntner (1902) in their important work on the Diplopoda of Madagascar. Originally, the genus 
contained a single species, Sphaeromimus musicus (DeSaussure and Zehntner, 1897, sub 
Sphaeropoeus). known only from a single male. Consequently, only male sexual characters were 
given with descriptive details focusing on the telopods. The unusual features of the species prompt- 
ed Jeekel (1999) to suggest that the then known Sphaeromimus specimen may have been "misla- 
belled or [represents] an introduced Indian sphaerotheriid". With the collection of male and female 
specimens of S. musicus at three different localities and the discovery of two new Sphaeromimus 
species, described below, it is now demonstrated that Sphaeromimus forms an established part of 
the endemic Malagasy fauna. Since the genus is no longer monotypic, genus-specific characters 
can be given. 

The genus Sphaeromimus can be distinguished from the only other Malagasy sphaerotheriid 
genus Zoosphaerium Pocock, 1895, on the basis of numerous characters. The genus description 
given below includes the characters DeSaussure and Zehntner (1902) mentioned in the original 
description of the genus. 

Diagnosis. — Members of the genus Sphaeromimus can be distinguished from Zoosphaerium 
by the following combination of characters: antennae short, with six joints, antennomeres without 
small spines and first antennomere without indentation. Apical antennomere rounded with numer- 
ous (up to 77) sensorial cones (apical antennomere cylindrical with four or more sensorial cones in 
Zoosphaerium), number of cones species-specific. Tarsi in Sphaeromimus broad (2.5-3 times 
longer than broad, Zoosphaerium up to 4.5 times longer than broad), tarsal tip densely covered with 
ventral spines. Anterior telopods with four joints {Zoosphaerium with three joints). Males with 
numerous strong stridulatory ridges on a plate termed 'harp' located on the first joint of the anteri- 
or telopods. Females with prominent, long stridulatory ridges on the subanal plate called 'wash- 
board.' Washboard divided into two parts by a suture of variable length depending on species. 
Cyphopod sclerites in the bursa of Sphaeromimus of unique shape. In Sphaeromimus, operculum 
of vulvae much longer than the 2"^^ leg coxa, without a central depression (operculum subreniform 
in Zoosphaerium). This high number of characters allows easy differentiation between the two 
Malagasy sphaerotheriid genera. 



562 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 29 

Description. — Known members of the genus range from 15 to 35 in body length, thoracic 
shield width ranges from 6.8 to 17.6. 

Head: only antennae with genus-specific characters, remaining features of head agree well 
with those found in most other sphaerotheriids. Eyes with numerous greenish ocelli, two of which 
are larger and one ocellus laterally displaced and separated (Figs. 24—25). Clypeus with single tooth 
(called labrum tooth by other authors), surrounded by hairs set in small pits. Tomosvary organ 
developed as a small round pit as in all known members of the order (Fig. 25). Center of posterior 
edge of head with or without patch of very small bristles (Figs 14, 39). 

Antennae: antennae short, six visible antennomeres more or less short and rounded. First 
antennomere without spines, 6* antennomere prominent, big, flat and longer than the others, car- 
rying many (40-77) sensorial cones (Figs 20, 43, 61). 

Mouth parts: external tooth of mandible with a distinct 'step' (Figs 18, 40), with 6 or 7 pecti- 
nate lamellae, apical teeth of pectinate lamellae broad and short (Figs 19, 41), number of teeth 
declining from apical to proximal pectinate lamellae. Gnathochilarium more or less hairy, with a 
few sensorial cones lateral of the palpi (Figs. 15-16, 47, 49). Centrally located pads (='Zapfchen- 
kappen' sensu Verhoeff) on the anterior edge of the lamellae linguales with sensorial cones (Fig. 
48). Tip of palpi with numerous sensorial cones distributed regularly around the tip. Epipharynx 
very similar in shape as known from other sphaerotheriid taxa (see Verhoeff 1928:841, fig. 419) 
(Fig. 45). 

Thoracic shield: ridges on lateral lobes of thoracic shield absent. Anterior rim of lateral lobes 
broad, used in volvation. 

Tergites: surface varies somewhat but mostly hairless and almost polished, except for the ante- 
rior paratergite depressions (see Material and Methods) which are more or less densely covered 
with hairs. Tergites always without a median keel. Tergites 3-12 each with a black carina ventral- 
ly on the anterior section of the tergites. Carinae apparently function as a locking device (Verhoeff 
1928:479), fitting over the rim of the lateral extension of the thoracic shield (Figs. 1, 28-29, 50). 
Endotergum variable, species-specific crenulations, marginal ridge and bristle patterns, marginal 
bristles branched (Figs. 17, 23, 44, 62). 

Sternite: first sternite with a sclerotized ledge along the anterior stemite lobe (Figs 4, 31, 53). 
Stemite lobe long, curved towards the legs, reaching the apical edge of coxa. Coxae and stemites 
without spines, but sternites three and beyond with a spine-like process which reaches about to the 
stigma opening of the anterior sternite. 

Anal shield: shape of anal shield not variable within genus. Males of S. musicus with a small 
invagination as described in other sphaerotheriids (VandenSpiegel et al. 2003; Jeekel 1986). Anal 
shield sometimes with a few small isolated hairs and a patch of hairs in the comers towards the 12* 
tergite. Ventral side of anal shield with single black locking carina (='Verschlussleiste' Verhoeff 
1928:479) on each side, locking carina with a slight central constriction (Figs. 3, 30, 52). 

Legs: remarkably short and broad, especially the tarsus, being only 2.5-3 times longer than 
broad. Tarsi of first two leg pairs with three to five ventral tarsal spines and a straight apical claw. 
Tarsi of leg pair 3-21 with 10-15 ventral spines on the apical part and a curved apical claw with 
one apical spine. Coxal lobes present, with small black triangular spines, variable in the genus. 
Femur with toothed ridge (Figs. 2, 26, 33, 51). Prefemur of last pair of legs basally with a small 
sclerotized knob on posterior side. 

Female sexual characters: subanal plate with washboard, consisting of well-developed stridu- 
lation ridges. Stridulation ridges always very long, ending just in front of the anterior margin of the 
washboard. Washboard with distinct median longitudinal groove, posterior rim of washboard with 
a central invagination. Shape of vulva unique. Operculum rounded and very long, always longer 



WESENER AND SIERWALD: REVISION OF THE GENUS SPHAEROMIMUS 563 



than the coxa and can reach about half of the length of the prefemur. Exterior and inner plates (EP, 
IP) below the operculum (termed bursa by Attems 1928; Jeekel 1974). Cyphopod sclerites consist- 
ing of two triangular apical sclerites and a much larger smoothly rounded third sclerite, all visible 
as dark structures near the suture of the vulva between inner and exterior plate (Figs. 5, 32). 

Male sexual characters: male gonopore conspicuous, located slightly above the middle and 
near the inside margin of the coxa of second pair of legs. Gonopores apparently complex, partial- 
ly closed by a round sclerotized plate carrying a few long hairs and featuring at least two membra- 
nous folds (Figs. 6, 27). Anterior telopods: with four joints in addition to the syncoxite. Harp on 
plate of first joint with three or more prominent stridulation ridges (Figs 8, 37, 58). Posterior side 
of second joint always with a large immovable lobe-hke flat projection. Lobe-like projection with 
some crenulation on the border juxtaposed the third and fourth joints. Fourth joint much thinner 
and longer than the proximal joints, about as long as the second and third joint combined. Apically 
with a single long sclerotized spine (spine A) on posterior surface, basally with two non-sclerotized 
spines (spines B). Spination sometimes variable within individuals, especially on the fourth joint. 
Distally with fringe of thick, long hairs (Figs. 9, 10, 38, 59). Posterior telopods: Syncoxite mesal- 
ly with lobe-like projections, termed inner horns (IH). Tips of inner horns (IH) with apical thorn 
and patch of hairs; terminal portion of inner horn bent posteriorly more than 90°. Subanal lobe 
densely covered with hairs (Figs. 11, 56). The 2"^ joint forms an immovable finger, the third joint 
forms a movable finger. Three characteristic non-sclerotized spines on the inside of immovable fin- 
ger, spaced at V3 intervals. Small triangular non-sclerotized lobe next to most proximal spine. Stout 
tip of immovable finger hook-shaped. Posterior face of movable finger with several sclerotized 
ridges. 

Variation. — Members of Sphaeromimus are small in comparison to Zoosphaeriiim, the lat- 
ter can reach a length of 100 mm (e.g., Z. hippocastanum), but moderate in size when compared to 
others in the order Sphaerotheriida. The number of stridulation ridges on the female washboard is 
correlated with the length of the individual, with three ridges on each side in the smallest females 
of Sphaeromimus splendidus sp. nov. and up to five ridges in the largest females. The number of 
ridges on each side of the washboard may vary in the same specimen. 

Natural history, behavior. — Life observations of the two newly described species 
revealed that the first pair of legs is not used when walking on flat ground. The first pair of legs is 
held up, above the ground and next to the head. Upon encountering an obstacle such as a leaf or 
twig (personal observations, senior author), the first pair of legs touches the obstacle. The first leg 
differs morphologically from the remaining legs by having fewer ventral spines and lacking the 
typical apical spine. The y'^-ll^^ leg pair show identical characters with litde variation, even in the 
same leg pair, regarding to the number of ventral spines and length of the claw. 

Living animals of S. splendidus sp. nov. and S. inexpectatus sp. nov. seem to avoid climbing 
on steeply inclined twigs. When lightly touched while on branches the animals quickly roll up and 
drop down. Haacker and Fuchs (1972) reported a different behavior from apparently arboreal 
species observed in South Africa: when touched while sitting on a branch, the animals coil up the 
head and anterior body, but hold on firmly to the branch using the posterior legs. Only after repeat- 
ed and aggravated disturbance the animals roll up and drop from the branch. One of the authors 
CT.W.) observed identical behavior as described by Haacker and Fuchs ( 1972) in one Zoosphaerium 
species found in Sainte Luce and Mandena, where they co-occur with the Sphaeromimus-species. 
This Zoosphaerium species was sometimes also found up to 250 cm high on trees and shrubs, feed- 
ing on the trunk. The behavioral differences may indicate different ecological niches for these sym- 
patric sphaerotheriid species. 

Discussion. — Currently, too few specimens are known to evaluate sexual dimorphism with 



564 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 29 

regards to the number of sensorial cones on the antennae as is known to occur in other sphaerotheri- 
id genera (Verhoeff 1928: 791). Regenerated antennae were observed in some specimens. In these, 
the number of sensorial cones was reduced. 

The black locking carinae on the inside of the anal shield show a central invagination in some 
specimens, which may indicate a fusion of originally two separate carinae. Verhoeff hypothesized 
that the anal shield of sphaerotheriids results from a fusion of at least two segments, the 13* seg- 
ment and the telson (Verhoeff 1928:448, Bitelotergit). The characteristics of the carinae described 
here represent further support for this notion. 

Species-specific characters found on the endotergum have also been reported from the South 
African genus Sphaerotherium (VandenSpiegel et al. 2003). 

The distribution of the here observed toothed ridge on the femora of all walking legs within 
the order is currently unknown, it may have been overlooked by other authors (Silvestri 1917: figs. 
5-10 and 17, Jeekel 1986, fig. 4). This ridge is present in all Malagasy sphaerotheriids examined to 
date by the senior author. Because of the rarity of female specimens, the vulvae were not dissect- 
ed. Thus, the exact form of the cyphopod sclerites cannot be illustrated here. 

The movable finger of the chela of the posterior telopods carries sclerotized ridges on its pos- 
terior surface. DeSaussure and Zehntner (1902) suggested these to represent another stridulation 
organ (Figs. 11-12, 35-36, 56-57). Haacker (1969:455) and VandenSpiegel et al. (2003) describe 
a similar feature in the South African Sphaerotherium and suggest that it may provide a better grip 
on the female legs during mating and we agree with this suggestion. The lobe-like projection on 
the 2"*^ joint of the anterior telopods with its small crenulations may serve a similar purpose. The 
function of the inner horns of the syncoxite of the posterior telopods is uncertain. It can be suggest- 
ed that the big spine on the inner horn of the syncoxite is used to open the female vulvae or to trans- 
fer the sperm, while the posterior and anterior telopods hold the female. Unfortunately, matings 
have been reported for only one sphaerotheriid species (Haacker 1968, 1969. 1974) who mentioned 
transfer of a spermatophore with the male legs. His observations appeared to indicate that females 
take the spermatophore into their mouths shortly after transfer of the spemathophor. However, dis- 
sections of the entire male and female head and SEM studies of the mouth parts revealed no spe- 
cial structure in the male mouth parts for sperm transfer and no visible sperm bag in the female's 
head. 

Conservation. — The two new species were found in two of the four remaining small patch- 
es of the southern littoral rainforest, in Mandena (1,103 ha, 160 ha slated as conservation area) and 
Sainte Luce (1,947 ha; Ramanamanjato et al. 2002, Vincelette et al. 2003). More field collecting in 
other areas may reveal other species of this interesting genus. Considering the fast destruction of 
the last isolated remaining forest patches (e.g.. Green and Sussman 1990) and the endemism of the 
here described new Sphaeromimus species in Madagascar, new studies in other areas of the island 
are urgently needed. 

Sphaeromimus musicus (DeSaussure and Zehntner, 1897) 

Figs 1 - 27 

Sphaeropoeus musicus DeSaussure and Zehntner, 1897 (publication of figure). 

SpJiaeromimus musicus, Saussure and Zehntner 1902 (publication of description). — Jeekel 1999 (lists species 
name) — Enghoff 2003 (lists species name). 

Material examined. — Type material: Male holotype; Madagascar, Province: unknown, coll. A. 
Grandidier. MNHN, CH038, vidi, without telopods, specimen figured in atlas published 1897, plate 4, figure 
1 a-e. Non-type material: 16 males, 3 females. Madagascar, Province: Toliara, coll. RNI Andohahela, par- 



WESENER AND SIERWALD: REVISION OF THE GENUS SPHAEROMIMUS 565 



eel II. camp 6. ~I20m NN, 24°49.0'S 46°36.6'E, 7-15.XII.1995, leg. S.Goodman, 2 males, 1 female; FMNH 
5378. 2 males; FMNH 5372. 1 male; FMNH 5409. 1 male, pitfall trap 16-18; FMNH 5407. Province: Toliara, 
1 male coll. Foret Analavelona, mid altitude forest with western and eastern elements, ~ 1050m NN, 9- 
15.III.1998. 22°40.7'S 44°11.5'E, leg. S. Goodman, 1 male; FMNH 5439. 2 males; FMNH 5427. Province: 
Toliara. coll. RP Berenty, Foret Bealoka, Mandrare River, gallery forest; ~35m NN, 24°57'25"S 46°16'17"E; 
3-8.II.2002. leg. B.L. Fisher et al., 5 males; 2 females, BLF 5315; CAS. 2 males, BLF 5314; CAS. 

Diagnosis. — Sphaeromimus musicus can be most easily distinguished from any other 
Malagasy sphaerotheriid by its unique coloration and pattern (Fig. 13), which identifies the species 
imambiguously. The body is orange, with an irregular black pattern near the posterior margin of 
each tergite. Each of the paratergites wears a median distinct thick black stripe. S. musicus is 
markedly more hairy than the other species of the genus, with hairs covering the head, legs, 
gnathochilarium, stemites and anal shield (Figs. 1-2, 4, 14-15). The anterior paratergite depres- 
sions carr}' an elongated patch of hairs on each. The body is less highly arched than in S. splen- 
didus sp. nov. The coxal lobes of the walking legs are only weakly developed, but somewhat big- 
ger than in 5. inexpectatus sp. nov. (Fig. 2). Remarkable is also the high number of over 75 sensu- 
al cones (Fig. 21) on the last antennomere, which is much higher than those of S. splendidus sp. 
nov. (Fig. 43). The female washboard (Fig. 7) and the male harp (Fig. 8) are the biggest known in 
all Malagasy sphaerotheriids, with the highest number of stridulation ridges in the genus 
Sphaeromimus. The shape of the female operculum is unique and shorter in S. musicus than in the 
other species of the genus. Its mesal margin is more strongly developed than in its congeners (Fig. 
5). The lower part of the inner plate (IP) of the female vulvae is not sclerotized and carries some 
triangular black spines. Molar plate process of the mandible with a single step (Fig. 17). In S. musi- 
cus, the endotergum features a distinct band of flattened nodules between the marginal bristles and 
the internal area covered with short spines and hairs (Fig. 17). 

Description. — Body length: 17.2-34.5; width of thoracic shield: 13.3-17.6; height of tho- 
racic shield 7.5-10.1. 

Habitus: In general, the tergites of this species seem to be higher than in most other Sphaero- 
theriida, with the exception of Sphaeromimus splendidus sp. nov. 

Coloration: body orange, with irregular black pattern near the posterior margin of each tergite. 
Each paratergite with a distinct thick black stripe, thoracic shield with even thicker black stripe, 
collum mostly black. Anal shield almost completely black (Fig. 13); head, antennae and legs 
orange-red. In alcohol, pattern and coloration, especially orange and red, are lost over time, either 
through exposure to light and/or alcohol; black stripe may fade completely, the black pattern 
becomes very irregular. The illustrated specimen in the original description shows this loss of col- 
oration clearly. For this study, we examined a number of specimens in different stages of coloration 
and pattern loss, from specimens featuring almost lifelike color and pattern to the stage illustrated 
by DeSaussure and Zehntner (1897). 

Head: with numerous hairs and setiferous pits mostly around the clypeus and lateral of the 
eyes. Some long, isolated hairs around the eyes and more distributed over the rest of the head. 
Posterior margin of the head towards the collum with dense field of very small hairs (Fig. 14). 

Antennae: shape as given in genus description. Length of antennomeres: 1>2>3=4=5<6; 6* 
antennomere being broadest and longest (Fig. 20), flat, reaching broadest point near the middle and 
does not taper towards the sensual plate, with up to over 75 sensual cones (Fig. 21). 

Mouth parts: mandible with six pectinate lamellae; number of teeth of pectinate lamellae 
declining from apical to proximal (Fig. 19). Molar plate process with a sharp single step near the 
apical border (Fig. 18j. Gnathochilarium ventrally with many hairs on the lamellae linguales. Field 
of four sensorial cones, three grouped together, the fourth displaced towards posterior margin, 



566 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 29 

located laterally of the palpi (Fig. 16). Epipharynx as in the genus description. 

Collum: anterior margin with two rows of isolated long hairs, posterior margin only with few 
isolated hairs, rows of hairs of the endotergum visible. 

Thoracic shield: with an area of numerous thick hairs on the concave lateral extension 
('Brustschildgruben' sensu Verhoeff) towards the marginal rim. Anterior rim of the lateral exten- 
sions broad (Fig. 1). 

Tergites: posterior margins of tergites three to seven with a visible fringe of short hairs, which 
originates from the endotergum. The anterior paratergite depressions of the tergites four to ten are 
densely covered with hairs, anterior paratergite depressions of the anterior tergites with several 
ridges each. Anterior paratergite depressions of tergites 11 and 12 also with pads of dense hair, but 
ridges not visible in intact specimens. Tips of posterior margins of paratergites project posteriorly. 
The endotergum features a distinct band of flattened nodules between the marginal ridge and the 
internal area covered with short spines and hairs (Fig. 17). 7^' Stemite: lobe long, reaching beyond 
the length of the coxa, covered with many long hairs and curved towards the leg pair (Fig. 4). The 
upper margin is smoothly rounded and completely covered with individual long hairs, lower mar- 
gin hairless (Fig. 4). 

Anal shield: rounded, neither bell-shaped nor tapered, in males there is a weak invagination 
not seen in females and less distinct than in the South African genus Sphaerotherium (Fig. 1:AI). 
The anal shield carries on both sides a black locking carina, sloping towards the posterior end of 
the anal shield (Fig. 3). The locking carinae in this species are well-developed, but narrow and of 
medium length compared to other species. 

Legs: tarsi of leg pair one and two with only four ventral spines and only weakly curved claws. 
Claws of the tarsi of following legs are curved wearing 12-14 ventral spines. Ninth pair of legs 
with a small lateral lobe and many small black triangular spines (Fig. 2). Coxae of all legs at the 
inside margin densely covered with many long hairs, also on the following leg joints at the inside 
margin some very long, isolated hairs. 

Female sexual characters: second pair of legs with coxal lobe. Operculum (Fig. 5:0) of vul- 
vae very broad and long, reaching Vs of the prefemur length. Mesal section of operculum drawn out 
apically and longer than lateral section. Center of operculum without indentation (=not subreni- 
form), lower margin straight. Exterior plate (Fig. 5:EP) of vulvae long and broad, its anterior mar- 
gin reaches around the base of the operculum. Inner plate (Fig. 5:IP) not as long as exterior plate, 
anterior margin of former ends below base of operculum. Posterior margin of inner plate not scle- 
rotized, sloping lower than exterior plate, with short, triangular black spines (Fig. 5). 

Subanal plate rounded, center of anterior margin with a broad shallow invagination. The wash- 
board with six strong, symmetrical stridulation ribs which end just in front of the anterior margin. 
Subanal plate divided by central suture not reaching anterior and posterior margins of subanal plate 
(Fig. 7). 

Male sexual characters: second pair of legs with a pronounced coxal lobe (Fig. 6). Anal shield 
with a weak invagination (Fig. 1:AS). Anterior telopods: first joint with a large stridulation harp 
and 5 stridulation ridges (Fig. 8). posterior side of second joint with a lobe-like projection, which 
reaches the 4'^ joint (Fig. 9). On its inside face two long, thin non-sclerotized spines (Figs. 
10:G-H). The outside face of the lobe carries a patch of very small (sensorial) hairs (Fig. 9:H). The 
third joint is short and slightly invaginated towards the lobe of the second joint. Near the invagina- 
tion insert two short (E) and one longer thin non-scletorized spine (F) (Figs. 9:E-F). The 4* joint 
carries basally a low knob (C) and a lateral non-sclerotized thin spine (D) (Figs. 9:C-D). The api- 
cal portion of the 2"^^ joint lobe is juxtaposed the low basal knob of the 4^^ joint (Figs. 9-10). 
Posterior telopods: telopod syncoxite densely covered with hairs. Outer surface of 2"'^ joint basal- 



WESENER AND SIERWALD: REVISION OF THE GENUS SPHAEROMIMUS 



567 



Table 1. Variation in S . musicus. * indi- 
cates specimens used for drawings and SEM; 
m: male; f: female; TS w: width of thoracic 
shield; SR: number of stridulation ridges of 
harp in males and washboard in females on 
left/right body side. 



ly with hairs, apically hairless. Stout tip of immovable finger hook-shaped. Chela without species- 
specific characters, movable finger with genus-specific dentition and row of crenulated teeth. The 
opposite finger (2"^ joint) features crenulations juxtaposed to the crenulated teeth of the movable 
finger. Base of movable finger laterally with some long hairs, more densely towards the outer mar- 
gin, apical section with a few sensorial hairs (Fig. 22). 

Distribution and ecology. — According to current collection records, S. musicus appears 
to be restricted to the southwestern region of Madagascar. To date, S. musicus is known from three 
localities, indicating a wider distribution range than some other sphaerotheriid species on 
Madagascar, e.g., S. inexpectatus sp. nov. and S. splendidus sp. nov. Sphaeromimus musicus was 
collected in gallery forests as well as in over 1,000 m elevation. It is remarkable that no specimens 
of this species were found in the spiny dry forest so widespread in its range, but the species appears 
to be restricted to semi-humid habitats such as gallery forests. The species was not found among 
other sphaerotheriid material, e.g., of the genus Zoosphaerium, collected in the eastern Hylaea 
areas or the w estem dry-deciduous forest. Collections 
took place during the wet season, December, 
Februar)' and the first half of March. Members of the 
genus Zoosphaerium were found at all three sites 
(Fig. 65) from which S. musicus were collected. No 
eggs were found in a dissected female collected dur- 
ing the wet season (Dec. 7-15, 1995 in RNI 
Andohahela, parcel II). It is unknown whether this 
species is active in the dry season. 

Conservation. — The currently fragmented dis- 
tribution of S. musicus is most likely the result of the 
continuing destruction of the natural vegetation. 
Habitat protection is vital for the survival of highly 
endemic species such as the type species of the genus 
Sphaeromimus. 

Discussion. — Males and females in Sphaero- 
theriida molt after maturity (pers. obser.). Onto- 
genetic changes of characters described above have 
not been investigated to date, e.g., it is possible that 
the number of stridulation ridges increases with the 
age and size of the animal. This seems to be the case 
in females, the number of stridulation ridges on the 
male harp remain constant (Table 1). The small size 
of the vulva in this species is remarkable when com- 
pared to the relatively larger female vulvae in the 
much smaller females of S. splendidus sp. nov. and S. 
inexpectatus sp. nov. 



Sex 


TSw 


SR 


Location 


m* 


17,0 


5/5 


RNI Andohahela, parcel 2 


m* 


16,0 


5/5 


RNI Andohahela, parcel 2 


f* 


14,5 


6/6 


RNI Andohahela, parcel 2 


m 


17,4 


5/5 


RNI Andohahela, parcel 2 


m 


15,8 


5/5 


RNI Andohahela, parcel 2 


m 


17,0 


5/5 


RNI Andohahela, parcel 2 


m 


16,5 


5/5 


RNI Andohahela, parcel 2 


m 


15,9 


5/5 


Foret Analavelona 


m 


15,4 


5/5 


Foret Analavelona 


m 


15,6 


5/5 


Foret Analavelona 


m 


17,4 


5/5 


RP Berenty 


m 


16,7 


5/5 


RP Berenty 


m 


16,7 


5/5 


RP Berenty 


m 


16,7 


5/5 


RP Berenty 


m 


13,3 


5/5 


RP Berenty 


f 


16,2 


8/8 


RP Berenty 


f 


15,5 


7/8 


RP Berenty 


m 


17,6 


5/5 


RP Berenty 


m 


16,5 


5/5 


RP Berenty 



Sphaeromimus splendidus sp. nov. 

Figs 28^9 



Material examined. — Type material: 1 female holotype; paratypes: 1 male; 3 females, 1 male 
immature, coll. Madagascar, Province: Toliara: Sainte Luce, littoral forest, 24°47'S 47°10'E; 08.IV.2003, leg. 
Wesener; FMNH 6702, 6703. 1 female (mature), identical collecting data; CAS. Other material examined: 
2 males (immature), 2 females (immature), coll. Madagascar, Province: Toliara: Sainte Luce, littoral forest; 



568 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 29 

24°47'S 47°10'E, 06.04.2003, leg. Wesener, 2 juvenile, (width of thoracic shield: 3.4 mm, 2.9 mm; body 
length 7.6 mm, 7.0 mm), same collection data; vouchers, deposited at the Universite Antananarivo. 

Diagnosis — S. splendidus is distinguishable from other Sphaeromimus species by its com- 
pletely black coloration and tergites with a satin sheen. This species is almost hairless, with only a 
few individual hairs on the anterior paratergite depressions and on the thoracic shield (Figs. 28-29). 
The body is more highly arched than in other Sphaeromimus-species. The coxal lobe is very long 
and well-developed which is one of the main characters by which this species can be distinguished 
from S. musicus and S. inexpectatus sp. nov. (Fig. 33). Remarkable is also the small number of only 
20^5 antennal cones and the 6* antennomere (Fig. 43) is very slender. The male anterior telopods 
differ from the telopods in S. musicus by possessing a small pointed process on the anterior side of 
the first joint, reaching the S^d joint. The operculum of the vulva reaches over the middle of the 
prefemur; its anterior margin is well rounded. The black locking carinae of the anal shield are short- 
er than in the other two species. The molar plate process of the mandible possesses one small and 
one big step (Fig. 40). The endotergum features only one row of marginal bristles and unique, 
rounded crenulations between the marginal ridge and the internal area, which is covered with short 
spines and hairs (Fig. 44). 

Description. — Body length up to 23.6; width of thoracic shield: 8.2-11.8 (mature); height of 
thoracic shield up to 6.5. 

Habitus: In general, the tergites of this species seem to be higher than in other Sphaerotheriida 
and higher than in all other species of this genus (Figs. 28-29). 

Coloration: The body is shiny black. Smaller specimens are creme-white with only a black 
posterior margin at each tergite. As the animals grow the black margins on the tergites expand until 
the tergites are completely black. Head and collum brown, antennae olive-blackish, but anten- 
nomeres five and six remarkably lighter in color. Legs also olive-blackish, but apically lighter in 
color. 

Head: with numerous hairs and setiferous pits mostly around the clypeus and lateral of the 
eyes. Few long, isolated hairs around the eyes and distributed over the rest of the head. Posterior 
margin of head towards the collum hairless (Fig. 39). Field of little crenulated teeth near the anten- 
nal socket with one small spine (Fig. 46). 

Antennae: shape as given in genus description. Length of antennomeres: 1>2>3=4=5<6. Sixth 
antennomere being longest (Fig. 42), flat, reaching broadest point near the middle, but is not broad- 
er than other antennomeres. Tapering only slightly towards sensorial disc. Only 20 to 45 sensorial 
cones (Fig. 43). 

Mouth parts: mandibular molar plate process with two steps near apical end (Fig. 40); with 
seven rows of pectinate lamellae, teeth short and broad; apical pectinate lamella with 18 teeth, 
number of teeth declining proximally (Fig. 41). Gnathochilarium ventrally with few hairs (Fig. 47), 
group of 4 sensorial cones located in a pit laterally of the palpi (Fig. 49). Epipharynx genus-like 
(Fig. 45). 

Collum: anterior margin with some isolated long hairs, posterior margin only with few isolat- 
ed hairs. 

Thoracic shield: with only very few small hairs on the concave lateral extension of the tho- 
racic shield towards the margin. Brim of anterior margin of lateral extension only slightly broader 
than remaining brim. 

Tergites: hairless, shiny, only the anterior paratergite depressions and their anterior margins 
with \'ery few short hairs. Anterior paratergite depressions of the anterior tergites with several 
ridges each. Anterior paratergite depressions of tergite 12 also with few hairs, but ridges not visi- 
ble in intact specimens. Tips of posterior margins of paratergites project posteriorly, stronger in ter- 



WESENER AND SIERWALD: REVISION OF THE GENUS SPHAEROMIMUS 569 



gites9-ll (Figs. 28-29). 

7" Stemite: lobe as long as coxa and curved to the leg pair. Upper margin smoothly rounded, 
isolated long hairs near the border. Rest of stemite hairless (Fig. 31: S = sternite). 

Anal shield: rounded, neither bell-shaped nor tapered (Figs. 22-23). Anal shield with distinct, 
broad, but short black locking carinae on each side, sloping towards the posterior end (Fig. 30). 
Remarkable is a very small invagination at the middle of the carinae. 

Legs: 9* leg pair with a pronounced coxal lobe and many small black triangular spines (Fig. 
33). Tarsi of first two leg pairs with only three to four ventral spines and only weakly curved claws. 
Tarsi of following leg pairs curved, with 10-14 ventral spines and the apical spine. Coxae mesally 
with many dense long hairs; other podomeres with few, very long, isolated hairs. 

Female sexual characters: second pair of legs with well-developed coxal lobe. Operculum (O) 
of vulvae: very broad and long, reaching over V2 of prefemur length; reaches its maximum length 
in center. Anterior margin without indentation (=not subreniform), lower margin with invagination 
in center. Exterior plate (EP) of vulvae long and broad, its anterior margin ends below base of oper- 
culum. Inner plate (IP) not as long as exterior plate, anterior margin of former extends below base 
of operculum (Fig. 32:0 = operculum, IP = inner plate, EP = exterior plate). 

Subanal plate rounded, center of anterior margin with a very broad shallow invagination. 
Washboard with three to five strong, symmetrical stridulation ribs, ending just in front of anterior 
margin. Washboard divided by central suture reaching anterior and posterior margins of subanal 
plate (Fig. 34) 

Male sexual characters: second pair of legs with coxal lobe. Anterior telopods: first joint with 
a small harp and three stridulation ridges (Fig. 37) and on its posterior side with a very small pro- 
jection, reaching the third joint. Posterior side of 2"^ joint with a lobe-like projection, reaching 4* 
joint (Fig. 38: A = big spine; B = two small spines). Third joint short, 4^^ as described in genus 
description (Figs. 37-38). Posterior telopods: telopod syncoxite nearly hairless. Chela without 
species-specific characters, movable finger with genus-specific dentition and row of crenulated 
teeth. Opposite finger (2"^^ joint) features crenulations juxtaposed the crenulated teeth of the mov- 
able finger. Base of movable finger laterally with some hairs. Stout tip of immovable finger hook- 
shaped (Figs. 35-36). Immature males with bud-shaped anlagen (primordia) in the place of 
telopods as in mature male. 

Distribution and ecology. — Some females collected in the beginning of April were carry- 
ing up to eight eggs, suggesting that the breeding season was in progress. Assuming a single annu- 
al breeding season and collecting adult egg-carrying females and juveniles with 19 leg pairs at the 
same time suggest that the adults are at least 2 years of age. 

So far this species of Sphaeromimus was collected only from a fragment of littoral rainforest 
on sand in Sainte Luce. This particular patch of littoral rainforest is virtually undisturbed and may 
represent the best preserved of all four still existing southern littoral forest patches (Dumetz 1999; 
Vincelette et al. 2003; deGouvenain and Silander 2003). Juveniles and adults could be found in 
thick (30-80 mm) leaf litter, containing mostly big leaves of trees. The leaf litter was wet and did 
contain also a large numbers of Spirostreptida, Isopoda, winged Blattodea, Diplura and 
Collembola. In this assemblage, the giant pill-millipedes were the biggest arthropods found. This 
species was found together with two species of the genus Zoosphaerium (description in progress) 
one of which occurs also in the littoral rainforest in Mandena and in the eastern lowland rainfor- 
est. The second Zoosphaerium species appears to be restricted to Sainte Luce. The forest patch of 
Mandena was intensively searched for 1 8 days without success for S. splendidus sp. nov. In addi- 
tion, S. splendidus sp. nov. was not found in any other collection samples. These observations sug- 
gest that S. splendidus sp. nov. is endemic to the littoral forest patch of Sainte Luce. Also, with 



570 



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Fourth Series, Volume 56, No. 29 



regards to the isopod fauna and vegetation (Dumetz 1999), the littoral forests of Mandena and 
Sainte Luce, albeit separated by a distance of only 20 km, display distinct faunal and floral differ- 
ences. Currently, both patches of littoral forests are separated by pseudosteppe with apparently lit- 
tle humus and soil arthropods (pers. observation). Maps showing forest distribution dating back to 
1950 indicate that continuous forest vegetation disappeared before 1950. Lehtinen et al. stated 
2003: ''At present, the landscape at Mandena and Sainte Luce is a series of littoral rainforest frag- 
ments in a matrix of extremely degraded anthropogenic sand-scrub. TJiis barren sand-scrub is the 



result of previous forest clearing, burning, and 
attempts at cattle grazing and is presumably a hos- 
tile environment for forest-dwelling organism (p. 
1359)." Our studies are comparable with this sug- 
gestion: no pill millipedes or other soil arthropods 
were found in the sand-scrub, no humus layer is vis- 
ible in the pseudosteppe. Actually, there are no geo- 
graphic barriers between the Mandena's and Sainte 
Luce's littoral rainforest, such as rivers and hills, 
which in other cases often form borders of a milli- 
pede species ranges. The only difference between 
the two localities is the annual precipitation, with 
higher rainfall in Sainte Luce (Donque 1972). 

Conservation. — The forest at Sainte Luce is 
subject to human impact and wood removal as one 
of us (T.W.) observed. Protecting this unique and 
still relatively pristine littoral forest should receive 
highest priority. 

Discussion. — Coloration not suitable for field 
identification, since shiny black Zoosphaerium 
species occur sympatrically. The only male known 
also shows the juvenile coloration, but has fully 
developed telopods and thus is most likely sexually 
active. 



Table 2. Variation in S. splendidus sp. nov. 
* indicates specimens used for drawings and 
SEM; m = male; f = female; TS w: width of tho- 
racic shield; SR: number of stridulation ridges of 
harp in males and washboard in females on 
left/right body side. * small, bud-shaped anlagen 
(primordia) of telopods present. 



Sex 


Status 


TSw 


SR 


m 


mature 


8.0 (4th segment!) 


3/3 


f (type) 


mature 


11,8 


5/5 


f 


mature 


11,2 


5/5 


f 


mature 


9,0 


4/5 


f 


mature 


8.4 


4/5 


f 


mature 


8,2 


4/5 


m 


immature 


5,1 


* 


m 


immature 


4,7 


* 


m 


immature 


3.1 


* 


f 


immature 


6,2 


4/4 


f 


immature 


5,1 


3/3 


7 


juvenil 


3,4 


- 


9 


juvenil 


2,9 


- 



Sphaeromimus inexpectatus sp. nov. 

Figs. 50-63. 

Type material. — l male holotype (width of thoracic shield: 7.3mm), 1 female paratype, in parts 
(w idth of thoracic shield: 6.8mm); Madagascar, Province: Toliara, Mandena; littoral forest; in leaf litter with 
small fruits. 24°57'l5"S 046°39'22"E ; IV.2003; leg. Wesener; FMNH 6701. 

Diagnosis. — Coloration unique in the genus, males of S. inexpectatus pink to red (Fig. 63). 
Species almost hairless, except for some isolated hairs on the anterior paratergite depressions and 
thoracic shield (Fig. 50). Sixth antennomere broader than in the other two Sphaeromimus species, 
with well over 70 antennal cones (Figs. 60-61). 

Coxal lobes only weakly developed. Lobe-like projection at the 2"'^ joint of the anterior 
telopods protruding laterally and reaching the distal end of the 3'^'^ joint (Fig. 58: F = one thin 
spine), a unique feature for this species. Sphaeromimus inexpectatus sp. nov. differs from S. musi- 
cus by the possession of a small process inserting on the anterior side of the first joint of the ante- 
rior telopods. extending to the 3'^'^ joint. Ver}' remarkable is the curs'ed. hook-like end of the immov- 
able finger of the posterior telopods (Figs. 56-57). The operculum of the vulva is large and extends 



WESENER AND SIERWALD: REVISION OF THE GENUS SPHAEROMIMUS 57 1 



over the middle of the prefemur. Its anterior margin is well rounded. The black locking carina of 
the anal shield is longer than in the other two species (Fig. 52: AS = anal shield; PL = pleurite). 
External tooth of the mandible with one big and a second small step. The endotergum features only 
one row of marginal bristles, which are separated by a wavy marginal ridge from the intermediate 
area covered with short spines and hairs (Fig. 62). 

Description. — Body length: circa 15; width of thoracic shield: 6.8 (f)-7.3; height of thoracic 
shield up to 4.5. 

Habitus: In general, the tergites of this species seem to be lower than in all other species of 
this genus. 

Coloration: body of mature male pink, posterior margin of each tergite with thin black line; 
immature female creme-white to reddish, posterior margin of each tergite with a broad, brown line. 
Head and collum in male type pink, anterior paratergite depressions gray to reddish; antennae and 
legs remarkably silver-gray to yellow. 

Head: with numerous hairs and setiferous pits mostly around the clypeus and lateral of the 
eyes. There are some long, isolated hairs around the eyes and more distributed over the rest of the 
head. The posterior margin of the head towards the collum is hairless. 

Antennae: shape as given in genus description; length of antennomeres: 1>2>3=4=5«6, last 
antennomere as long as antennomeres 4+5 combined; last Antennomere flat and very broad (Figs. 
60-61). 

Mouth parts: mandibular molar plate process with one big and a second smaller step near the 
apical tip; with seven pectinate lamellae, 20 teeth in apical pectinate lamella, number declining 
proximally. 

Collum: anterior margin with some isolated long hairs, posterior margin only with few isolat- 
ed hairs. 

Tlwracic shield: with only very few short hairs on the concave lateral extension of the thoracic 
shield tow ards the marginal rim. Rim around anterior margin only slightly broader than around the 
rest of the thoracic shield. 

Tergites: hairless with very few short hairs in the anterior paratergite depressions and with 
some more longer hairs on the anterior margin. Tips of posterior margins of paratergites do not 
project posteriorly (Fig. 50). 

1st Stemite: lobe as long as coxae, with some isolated long hairs, curved towards leg pair, 
upper margin irregularly rounded with two invaginations (Fig. 53: S = sternite), a few isolated long 
hairs near the margin. Rest of the stemite hairless. 

Anal shield: rounded, neither bell-shaped nor tapered. Anal shield with black locking carinae 
on each side, sloping towards the posterior end of the anal shield (Fig. 52: AS = Anal shield; PL = 
pleuritej. The locking carinae in this species are well-developed and broad, remarkably longer than 
those of the other Sphaeromimus species. Locking carinae with distinct but very small invagination 
at the center. 

Legs: the first leg pair with only three, the 2"^ with four to five ventral spines and only weak- 
ly curved claws. Claws of the following leg pairs are curved. Coxal lobe at 9* leg pair very weak- 
ly developed, with many small black triangular spines (Fig. 51). Tarsi of remaining legs with 12- 
15 ventral spines and one apical spine (damaged in specimen). Coxae at mesal margin with many 
dense long hairs, also on following podomeres some very long, isolated hairs. 

Female sexual characters: 2"^ pair of legs without coxal lobe but with one black spine. 
Operculum (O) of vulvae very broad and long, reaching V2 of the prefemur length, maximum length 
in the center. Center of operculum rim without indentation (=not subreniform), lower margin with 
weak invagination in the center. Exterior plate (EP) of vulvae long and broad, its anterior margin 



572 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 29 

ends below the base of the operculum. Inner plate (IP) is not as long as exterior plate, anterior mar- 
gin of former ends below base. (Fig. 54: O = operculum, EP = exterior plate, IP = inner plate) 

Subanal plate rounded, center of anterior margin with a broad invagination. The washboard 
with three strong, symmetrical stridulation ribs, ending just in front of the anterior margin, pt and 
S^d ribs smaller than 2"^ . Subanal plate divided by short median suture only in the center (Fig. 55). 

Male sexual characters: 2"*^ pair of legs without a coxal lobe. Anterior telopods: l^t joint with 
small harp and three stridulation ridges (Fig. 58: F = one thin spine), posterior side of l^t joint with 
a small projection which reaches the S'''! joint. Lobe-like projection laterally on posterior side of 2"^ 
joint reaching 4* joint (Figs. 58-59). Si'd joint short and slightly invaginated towards the lobe of the 
second joint, with one longer thin non-scletorized spine (F) juxtaposed the second joint lobe (Figs. 
58-59: A = big spine, B = two small spines, D = small lateral spine. F = longer spine). Posterior 
telopods: Movable finger of chela with genus-specific dentition and row of crenulated teeth. The 
opposite finger {2^'^ joint) features crenulations juxtaposed to the crenulated teeth of the movable 
finger and also one non-slerotized spine on its anterior side (Fig. 56). Movable finger almost hair- 
less. 2"d joint with some hairs on the immovable finger. Stout tip of immovable finger curved and 
hook-shaped (Figs. 56-57). Telopod coxa densely covered with hairs. 

Distribution and ecology — So far this species of Sphaeromimus was collected only from 
a fragment of littoral rainforest on sand in Mandena. This particular patch of littoral rainforest is 
little disturbed, with 50-75% forest cover (QIT Madagascar forest map). The holotype was found 
in thin (5-30mm) dry leaf litter, containing mostly leaves and some tree fruits. A few winged 
Blattodea were found as well. A new species of genus Zoosphaeriiim (unpublished/in preparation), 
which occurs also in the littoral rainforest in Sainte Luce and in the eastern lowland rainforest, was 
common in this area (>300 mature and mostly immature where detected). 

Body rings of spirostreptid and small sphaerotheriid tergites were found in a layer of arthro- 
pod remains around ant holes of a big red ant species. It is unknown how the ants are able to hunt 
these well-armored animals. Rolled up sphaerotheriids were placed near ants, but the ants showed 
no interest. A large Zoosphaerium specimen (34 mm long. 16 mm broad (2"^ segment)) was put in 
a cage with one Carnivora: Galidia elegans inside, which was caught and kept at the Pepiniere in 
Mandena. Galidia was able to detect the rolled up specimen, broke the tergites with a few bites of 
the lateral teeth and ate internal parts, ignoring the intestine tergite pieces. It is likely that Galidia 
elegans may represent a predator of pill millipedes, including Sphaeromimus inexpectatus. 
Predation of pill millipedes by mongoose was reported by Eisner and David (1969). 

The female collected in the middle of April was carrying two eggs, suggesting that the breed- 
ing season was in progress. The forest patch of Mandena was intensively searched for 18 days dur- 
ing rain and at night without locating more specimens. This fact prompts us to suggest that S. inex- 
pectatus is either a very rare species or was not active during the collection time. S. inexpectatus 
was not found in any other collection samples or in nearby littoral forest patches of Petriky and 
Sainte Luce. Additionaly it was not present in the collections of CAS. FMNH or in the huge col- 
lections of the MNHN. People living in the area are familiar with pill millipedes, calling them 
"Mia." but were not aware of this red-colored species. These observations may indicate that S. 
inexpectatus is endemic or now restricted to the littoral forest patch of Mandena. According to the 
isopod fauna and vegetation (Dumetz 1999), the littoral forests of Mandena are different from those 
of Petriky and Sainte Luce, albeit a distance of only 20-30 km separates these. 

Conservation. — Currently, the observed patch of littoral forests is separated by pseu- 
dosteppe or Eucalyptus plantations with apparently little humus and soil arthropods (per. observa- 
tion). In the past 50 years almost 137c of the original forest was destroyed (Vincelette et al. 2003). 
CuiTently. the small study area is efficiently protected by QIT Madagascar. It is however, uncer- 



WESENER AND SIERWALD: REVISION OF THE GENUS SPHAEROMIMUS 



573 



Table 3. Species separation in Sphaeromimiis. No. = Number; gn 
telopods: p.t. = posterior telopods; SR = stridulation ridges. 



gnathochilarium; a.t. = anterior 



Character: 



Sphaeromimus musicus 



Sphaeromimus splendidus Sphaeromimus inexpectatus 
sp. nov. sp. nov. 



Tergite coloration: 


orange with black pattern 


black 


Body length: 


up to 34.5 mm 


up to 23.6 mm 


No. of SR in female 


7-Aug 


5-May 


No. of SR in male 


5 


3 


antennal cones 


up to 75 


up to 45 


Sensorial cones lateral of 


4. 1 displaced 


A, all together 


gn-palpi 






Surface of tergites 


few hairs 


bald 


Patch of hairs on the head 


present 


absent 


towards the collum 






Molar plate process of 


with 1 large step 


with 2 steps 


mandible 






No. of pectinate lamellae 


6 


7-Jul 


No. of oceUi 


>80 


50-60 


Coxal lobe of legs 


weakly developed 


strongly developed 


Endotergum: marginal 


straight 


straight 


ridge 






Endotergum: flattened 


oval 


rounded 


nodules 






Endotergum: No. of rows 


3-Mar 


1-2 


of marginal bristles 






a.T process of 2nd joint 


only posterior of joint 3&4 


only posterior of joint 2 


visible 






p.t.: 2nd joint 


apical end stout 


apical end stout 



pink 

at least 15 mm 

3 

3 

up to 75 

9 

bald 
absent 

with 2 steps (2nd small) 

7 
? 

nearly absent 
curved 

rounded 
1 



apical end hook-like 



tain, whether the protected area is large enough to sustain viable populations of this species. The 
senior author noted the lack of old large trees in the area and wood removal by humans is ongoing. 



Discussion 

The three species of Sphaeromimus are easily distinguished from each other, see Table 3. 
Jeekel (1974, Fig. 64 B) presented the most recent classification of the order Sphaerotheriida, while 
Hoffman (1976) and Mauries (2001) modified Jeekel's classification of the sphaerotheriid family 
Sphaeropoeidae (Fig. 64 A). Jeekel employed characters found in the shape of the female vulva and 
the stridulation organs (harp in males, washboard in females) to separate tribes and subfamilies. 
The genus Sphaeromimus belongs to the family Sphaerotheriidae, sharing the main synapomorphy 
of its genera: basis of the vulval operculum embraced by the bursa (consisting of the exterior and 
inner plate. Fig. 64, character 1). Jeekel considers the presence of a female stridulation organ, the 
washboard (Fig. 64, character 2) as the synapomorphy for the subfamily Arthrosphaerinae, to 
which the genus Sphaeromimus is currently assigned. The other synapomorphy of the subfamily 
cited by Jeekel, the median protrusion of the bursa, is not present in Sphaeromimus (Fig. 64, char- 
acter 3j. In Jeekel's classification, Zoosphaerium and Sphaeromimus, the two Malagasy 
sphaerotheriid genera, are placed in the tribe Zoosphaeriini, based on the possession of the harp in 
the males (occurs in both genera, character 4). Jeekel also listed the shape of the subreniform 



574 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 29 

female operculum (character 5), as it occurs in Zoosphaerium, as an apomorphy of the tribe. 
However, now that females of Sphaeromimus are known, this latter apomorphy cannot be support- 
ed. Females of Sphaeromimus have a round operculum with a smooth edge. 

Furthermore, Sphaeromimus shares characters with members of the Indian genus 
Arthrosphaera, currently placed by Jeekel (1974) in the tribe Arthrosphaerini of the 
Arthrosphaerinae. Such shared characters are: 6* antennomere flat and broad (cylindrical in 
Zoosphaerium, Fig. 64, character 8), and the four-jointed anterior telopods (Attems 1936, Fig. 64, 
character 9). Thus, males of Sphaeromimus share on the one hand a characters with the genus 
Arthrosphaera (characters 8 and 9) and on the other hand a character, the harp (character 4), with 
the genus Zoosphaerium (DeSaussure and Zehntner 1902; Pocock 1895). In Sphaeromimus, the 
washboard features a rather deep median groove (character 10) of variable length. The presence of 
the groove may indicate the fusion of two separate plates. In contrast, all Zoosphaerium species 
examined to date possess a completely fused subanal plate without a suture or groove. A groove is 
also present in at least one species of the Indian genus Arthrosphaera. Unfortunately, the form of 
the subanal plates are known for only few members of both genera. These morphological details 
indicate clearly that the current classification scheme (Fig. 64) lacks sufficient character support 
and that more characters are needed to define monophyletic clades within the Sphaerotheriida 
unequivocally. 

Acknowledgments 

The senior author gratefully acknowledges the support and advice by Prof. Dr. J.-W. Wagele. 
We thank the Direction des Eaux et Forets and the Commission Tripartite for their authorization to 
carry out this work. QIT Madagascar Minerals and their environmental and conservation team 
headed by Manon Vincelette and Jean-Baptiste Ramanamanjato provided excellent support. The 
paper is part of the Accords de Collaboration between the Universite d' Antananarivo (Dept. 
Biologie Animale and Anthropologic et Biologic Evolutive), QIT Madagascar Minerals and 
Hamburg University. The support by O. Ramilijaona and D. Rakotondravony is gratefully 
acknowledged. During fieldwork QIT Madagascar provided logistics support (arranged by J.B. 
Ramanamanjato). Collecting pemiits were arranged by Dr. O. Ramilijaona (Department de Biologie 
Animale, Universite d' Antananarivo). Special thanks go to the Direction des Eaux et Forets, 
Antananarivo for arranging collecting and export permits. The senior author conducted the field- 
work on an expedition organized by Prof. Dr. J.U. Ganzhorn (University of Hamburg). Expenses 
for the fieldwork were defrayed through personal funds. We thank Mr. Charles E. Griswold and Mr. 
Darrell Ubick (California Academy of Sciences) for the loan of material. Special thanks go to Dr. 
C. Schmidt (Ruhr-University Bochum) for continuous advice throughout the study. Ms B. Strack 
(Field Museum) assisted us during the preparation of the SEM images. We thank Drs. 
VandenSpiegel. Hamer and Ganzhorn for helpful suggestions on previous drafts of the manuscript. 
A one-month internship of the senior author at the Field Museum of Natural History and the work 
on the manuscript was supported by NSF grant DEB 97-12438 to P. Sierwald and W.A. Shear 
(Hampden-Sydney College. Virginia). A 20-day visit of the senior author at the Museum National 
d'Histoire Naturelle to study the type collection and the undetermined material was supported by 
Synthesys grant FR-TAF-186. Special thanks go to J. -J. Geoffroy and L.E. Leoz for the organisa- 
tion of this visit. 

Literature cited 

Attems. C. 1897. Myriopoden. Abhandhmgen herausgegeben von der Senckenbergischen Naturforschenden 
Gesellschaft 23:473-536 + 4 Tafeln 



WESENER AND SIERWALD: REVISION OF THE GENUS SPHAEROMIMUS 575 



Attems, C. 1928. The Myriopoda of South Africa. Annals of the South African Museum 26:213-240. 

Attems, C. 1936. Diplopoda of India. Memoirs of the Indian Museum 11:136-195 

Attems, C. 1942. Neue Sphaerotheriden des Wiener Museums. Annalen des Naturhistorischen Museums in 

Wien 53 11:60-73; plates VII-IX. 
CR-AWFORD, C. S. 1992. Millipedes as model detritivores. Berichte des Naturwissenschaftlich-Medizinischen 

Verein Innsbruck 10:277-288. 
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Biogeography and Ecology in Madagascar. Monographiae Biologicae No. 21. The Hague, Netherlands. 
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the world's hottest hotspots on its way out. Oryx 35(4):346-348. 
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Druck, Leverkusen, Germany. 480 pp. 
Green, G.M., ant? R.W. Sussman. 1990. Deforestation history of the eastern rain forests of Madagascar from 

satellite images. Science 248: 212-215. 
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Haacker, U. 1969. Das Sexualverhalten von Sphaerotherium dorsale (Myriapoda, Diplopoda). Zoologischer 

Anzeiger, Supplement, (Verhandlungen der Deutschen Zoologischen Gesellschaft) 32:454-463. 
Haacker. U. 1974. Patterns of communication in courtship and mating behaviour of millipedes (Diplopoda). 

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pedes (Diplopoda). African Entomology 10; 1; 29^2 
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Jenkins, P.D. 1993. A new species of Microgale (Insectivora:Tenrecidae) from eastern Madagascar with an 

unusual dentition. American Museum Novitates (3067): 1-11. 
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veau Prionobelum (Diplopoda, Sphaerotheriida. Sphaeropoeidae) de Hainan, Chine. Zoosystema 23(1): 

131-142. 
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Tasforests 10:141-157 
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Akademische Verlagsgesellschaft. Leipzig. Germany. 
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Inc.. New York. New York, USA. 



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ILLUSTRATIONS 



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Coxa 



Figures 1^. Sphaeromimus musicus, male. 1: habitus; 2: left 9th leg, posterior view; 3: anal shield, dorsal view of black 
locking carinae; 4: 1st right stemite with coxa of 1st pair of legs. AI = invagination of anal shield; 12T = 12th tergite; PL = 
pleurite: .AS = anal shield: S = sternite. Scale bars: 1 mm. 



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579 




5&7 




Figures 5-7. Sphaeromimus musicus, female and male. 5: 2nd left leg: coxa (female) with vulva; 6: 2nd left leg: coxa 
Cmale;, posterior view; 7: washboard. O = operculum; IP = inner plate; EP = exterior plate. Scale bars: 1 mm. 



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Figure 13 (above). Sphaewmimus musicus. Photo of freshly preserved male. 



Figures 8-12 (left). Sphaewmimus musicus, male. 8: left anterior telopod, anterior view; 9: left anterior telopod. pos- 
terior view; 10: anterior telopod, lateral view; 11: posterior teiopods, anterior view; 12: posterior right telopod. posterior 
view. A = 4th joint big spine; B = 4th joint 2 small spines; C = 4th joint knob; D = 4th joint 1 small lateral spine; I - 2nd 
joint sensorial hairs; G = 2nd joint lobe crenulation; H = 2nd joint lobe 2 spines; IH = inner horns. Scale bars: 1 mm. 



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Figures 14-19. Sphaeroniimus miisicus. male SEM. 14: patch of hairs on head to collum: 15: gnathochilarium, ventral 
view: 16: field of sensorial cones lateral of palpi of gnathochilarium: 17: endotergum 18: right mandible, general view; 19: 
right mandible, pectinate lamellae. 



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Figure 20. Sphaeromimus musicus, male SEM, antennae, lateral view. 



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Figures 21-25. Sphaewmimus musicus. male SEM. 21: 6th joint of antennae; 22: apical part of movable finger of pos- 
terior right telopod: 23: bristle of endotergum: 24: right ocelli: 25. antennal groove with Tomosvary organ (TO) and aber- 
rant ocellus. 



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Figures 26-27. Sphaewmimus musicus, male SEM. 26: posterior side of 9th femur with toothed ridge; 27: 2nd coxa. 
posterior view, coxa] lobe and male gonopode. 



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28&29 




Figures 28-31. Sphaewmimus splendidus. female holotype. 28: habitus, right side; 29: habitus, left side; 30: anal 
shield, dorsal view of black locking carinae; 31: 1st right stemite; 12T = 12th tergite; AS = anal shield: PL = pleurite; S = 
stemite. Scale bars: 1 mm. 



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Figures 32-34. Sphaeromimus splendidus, female holotype. 32: left vulva; 33: left 9th leg, posterior view; 34: wash- 
board; O = operculum; IP = inner plate; EP - exterior plate. Scale bar: 1 mm. 



588 



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syncoxite 



syncoxite 



Figures 35-38. Sphaeromimus splendidiis, male paratype. 35: left posterior telopod. anterior view; 36: left posterior 
telopod, posterior view; 37: left anterior telopod. anterior view; 38: left anterior telopod, posterior view; A = big spine; B = 
two small spines; IH = inner horns. Scale bar: 1 mm. 



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Figures 39^1. Sphaewmimus splendidus, female SEM. 39: patch of hairs on head lo collum; 40: right mandible, molar 
plate process; 41: right mandible, pectinate lamellae. 



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FiGL'RES 42-44. Sphaeromimus splendidus. female SEM. 42: antennae lateral: 43: 6th antennomere; 44: endotergum. 



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Figures 45-46. Sphaewmimus splendidus, female SEM. 45: epipharynx, anterior side; 
with crenulated teeth and one spine. 



edges of antennal groove 



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Figures 47-49. Sphaeromimus splendidus. female SEM gnathochilarium. 47: gnathochilarium, ventral view; 48: sen- 
sual cones on medial pads: 49: pit laterally of palpi with sensual cones. 



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53 




Figures 50-53. Sphaewmimus inexpectatus, male holotype. 50: habitus; 51: left 9th leg, posterior view; 52: anal shield, 
dorsal view of black locking carinae; 53: left 1st stemite; AS = anal shield, PL - pleurite; S = stemite. Scale bars: 1 mm. 



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Figures 54-55. Sphaew- 
mimus inexpectatus, female 
paratype; 54: vulva (macerat- 
ed); 55: washboard right; O = 
operculum; IP = inner plate; 
EP = exterior plate. Scale bars: 
0.5 and 1 mm. 




54 = 0.5 mm 



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595 




Figures 56-59: Sphaeromimus inexpectatus. male holotype; 56, left posterior telopod, posterior view; 57, posterior 
telopods, anterior view; 58, anterior telopods, anterior view; 59, anterior telopods, posterior view; A = big spine; B = two 
small spines; D = small lateral spine; F = longer spine; IH = inner horns of posterior telopod. Scale bar: 1 mm. 



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Figures 60-62. Sphaewmimus inexpectatus. male holotype SEM. 60: antennae lateral; 61: 6th antennomere; 62: 
endoter2um. 



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Figure 63. Sphaewmimus inexpectatus. male holotype, photo. 



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A 



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Sphaerotheriidae 



Sphaerotheriinae 



Arthrosphaerini 



Arthrosphaerinae 



Zoosphaeriini 



"Sphaeropoeidae" 



Sphaeropoeinae 



Sphaerotheriini South Africa 



Cyliosomatini Australia, New Zealand 



India 



Madagascar 



Arthrosphaera 
other Arthrosphaerini 
Zoosphaerium 
Sphaeromimus 

Sphaeropoeini Southeast Asia 

Indosphaerini India 

Castanotheriini Malaysia, Philippines 

Rajasphaerinae Borneo 



B 



Sphaerotheriinae 



Sphaerotheriidae 



(2), (3) 



(6) 

Arthrosphaerini 



Arthrosphaerinae 



2,3 



(4) 



Zoosphaeriini 



"Sphaeropoeidae" 



(1) 



(7) 



Sphaerotheriini South Africa 



Cyliosomatini Australia, New Zealand 

Arthrosphaera 
8,9,10,11 



India 



other Arthrosphaerini 



Zoosphaerium 



5, (8), (9), (10), (11) Madagascar 



Sphaeromimus 



(5), 8, 9, 10,11 
^ Sphaeropoeinae Southeast Asia, incl. India 



Castanotheriini Malaysia, Philippines 



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599 




Figure 65. Distribution map of Sphaeromimus. 



Figure 64 (left). Classification of the order Sphaerotheriida translated into a cladogram, with geographical distributions 
of clades. A. After Hoffman 1976, 1980, with modifications by Mauries 2001 incorporated. B. After Jeekel 1974. Shelley 
(2003) recommended use of the family name Zephronidae instead of "Sphaeropoeidae." Numbers 1-7 on branches of Fig. 
64 B indicate group-defining characters used by Jeekel (1974), numbers in parentheses indicate absence of character; l=vul- 
val operculum embraced by bursa, 2=female washboard present, 3= medium protrusion of bursa, 4= male harp present at 
anterior telopod. 5= vulval operculum subreniform, 6= stridulation organ on posterior male telopods, 7= movable digit of 
posterior telopod consists of two distinct podomeres, (7)= movable digit of posterior telopod consists of single podomere. 
Numbers 8-10 indicate characters and their distributions discussed in this study: 8= 6th antennomere flat and broad. (8)=6th 
antennomere cylindrical. 9= four-jointed anterior telopod in males, 10= female washboard divided (known from only a sin- 
gle Arthrosphaera species), (10)= female washboard undivided (with other variable features in Zoosphaerium). 11= oper- 
culum well rounded, (11)= operculum with central depression. 



Copyright © 2005 by the California Academy of Sciences 
San Francisco, California, U.S.A. 



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Fourth Series 

Volume 56. No. 30. pp. 601-605. 4 figs. December 30, 2005 



A Remarkable New Species of Acropyga 

(Hymenoptera: Formicidae) from Gabon, 

with a Key to the Afrotropical Species 

John S. LaPoUai and Brian L. Fisher^ 

1 Depaitinent of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, 

D.C. 20013-7012 USA; Email: lapollaj@si.edu: - Department of Entomology, California Academy of 

Sciences, San Francisco, California, 94103, USA; Email: bfisher® calacademy.org 

A new species o^ Acropyga, A. bakwele sp. nov., is described from Gabon. This is an 
intriguing species because unlilie any other known because its worker possesses a 
median ocellus, unlike any other known Acropyga. This species is the largest 
Acropyga known from Africa, and one of the largest in the world. In overall appear- 
ance, the worker resembles the southern African A. arnoldi. A key to Afrotropical 
Acropyga is provided. 

Ke\'words: Acropyga, Afrotropical, Formicinae, Gabon, Hymenoptera, Lasiini, trophophoresy. 

Acropyga are small formicine ants known for their habit of tending mealybugs underground 
on plant roots (Bunzli 1935; Weber 1944; Johnson et al. 2001; LaPolla et al. 2002). The relation- 
ship between the ants and mealybugs is complex (LaPolla et al. 2002; LaPolla 2004), and perhaps 
the most spectacular expression of this complexity is the fact that virgin queens emerge from their 
birth nests carrying a mealybug between their mandibles to presumably serve as a seed individual 
for the new ant colony. This behavior has been termed trophophoresy (LaPolla et al. 2002). 

A recent world revision of Acropyga revealed 37 species (LaPolla 2004). One of the surpris- 
ing results of that study was the lack of Acropyga species diversity from the rainforests of West and 
Central Africa. In other rainforest areas, such as in Southeast Asia and the Nootropics, Acropyga 
species diversity surpasses at least a dozen species in each region. LaPolla (2004) reported only 
two species from Africa, A. arnoldi and A. silvestrii. It remained unclear if Africa simply possessed 
a lower number of Acropyga species for unknown reasons (there are now a total of three species 
known from the continent), but the relatively few collections from western and central Africa may 
be indicative of a collecting artifact. In support of a collecting bias giving a lower number of 
Acropyga species than actually present in West and Central Africa, we report here on a new, inter- 
esting species recently collected in Gabon. Given the recent world revision by LaPolla (2004), we 
were able to recognize this new species and place it within a comparative framework. 

Materials and Methods 

In February 1998. BLF participated in a biological inventory of the Minkebe forest, an area of 
about 32.000 km^, in northeastern Gabon. Goodman et al. (2001) provides additional details on the 
inventory. The Minkebe forest is composed of a large block of Guineo-Congo lowland forest that 
drains a vast area. The northern area of that forest is part of the Ntem River watershed and the rest 
enters into the Ivindo River. The inventory was near the northwestern boundary of the Minkebe 

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Protected Area in an area of pockets of mixed heterogeneous and Maranthaceae forests within a 
vast area of marshland. This region is part of the Aya River drainage, which forms one of the main 
tributaries of the Ntem River. Our camp was in place between 5 and 17 February 1998 and was 
located in the Province de Woleu-Ntem, 28 km ESE Minvoul 2°05.2'N, 12°22.5'E, 600 m a.s.l. We 
began our trek into the forest from the Baka village of Doumasi, along the Ntem and to the east of 
Minvoul. Three distinct habitats types were found adjacent to the camp: marshlands dominated by 
Raphia, heterogeneous forest, and homogeneous forest composed of Gilbertiodendron. The leaf lit- 
ter transect that collected the Acropyga described here (BLF1684) was from forest adjacent to the 
marsh. The soil was moist and sandy. 

All measurements were taken at 80 x power with a Leica MZ 12 microscope using an orthog- 
onal pair of micrometers and recorded to the nearest 0.001mm and rounded to two decimal places 
for presentation. All measurements are given in millimeters. Digital images (Figs. 1-4) were cre- 
ated using a JVC KY-F75 digital camera and Syncroscopy Auto-Montage (v 5.0) software. 
Morphological terminology employed throughout follows Bolton (1994), with modifications 
where noted. Anatomical abbreviations are elaborated here: 

TL: Total Length: HL+ML+GL. 

HL: Head length: the length of the head proper, excluding the mandibles; measured in full-face view from 
the midpoint of the anterior clypeal margin to a line drawn across the posterior margin from its highest 
points (to accommodate species where the posterior margin is concave). 

HW: Head Width: the maximum width of the head in full-face view (excluding the portion of the eyes that 
extend pass the lateral sides of the head). 

SL: Scape Length: the maximum straight line of the antennal scape excluding the condylar bulb. 

ML: Mesosoma Length: the length of the mesosoma (=alitrunk) in lateral view from the anterior most point 
of the pronotum (including the "neck" of the pronotum) to the posteriormost point of the metapleuron. 

GL: Gaster Length: the length of the gaster in lateral view from the anteriormost point of first gastral seg- 
ment (third abdominal segment) to the posteriormost point of the acidopore. 

CI: Cephalic Index: HW = 100/HL. 

SI: Scape Index: SL = 100/HW. 

Systematic Treatment 

Acropyga bakwele LaPolla and Fisher, sp. nov. 

Figures 1-4. 

HOLOTYPE WORKER. — Gabon: Province Woleu-Ntem, 31.3 km 108° ESE Minvoul, 2°04.8'N, 
12°24.4'E, elev. 600 m ll.ii.l998, sifted leaf litter, rainforest (coll. B.L. Fisher) collection code: BLF01684, 
specimen code: CASENT0104123 (CASC) 

Diagnosis. — 8-toothed mandible: mandibular apical tooth about twice as along as other teeth; 
median ocellus present; total length > 3 mm. 

Description. — Worker; Overall appearance similar to Acropyga arnoldi and A. silvestrii, 
see LaPolla (2004) for details of these two species. Head (see Fig. 2); reddish-yellow; head slight- 
ly longer than wide; covered in a thick layer of appressed hairs, with short erect hairs along poste- 
rior margin; posterior margin slightly concave medially; median ocellus present; eyes relatively 
large for an Acropyga (ca. 10 facets) and placed at lower Va of head; 11-segmented, incrassate 
antennae; scape surpasses posterior margin by about half length of pedicel; scape with thick layer 
of appressed hairs, scattered erect hairs throughout; clypeus slightly convex medially; mandible 

Figures 1-4 (right). Acropyga bakwele. sp. nov. 1) lateral view: 2) head in full-frontal view: 3) mandible in full frontal 
view; 4) dorsal view. 



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broad, with six distinct teeth; mandibular basal angle distinct, but not forming seventh tooth; api- 
cal tooth twice as long as other teeth (Fig. 3). Mesosoma (see Figs. 1, 4): reddish-yellow; in later- 
al view, pronotum with short anterior shelf; dorsum covered in layer of appressed hairs, with scat- 
tered erect to suberect hairs throughout; metanotal area distinct; propodeum rounded; declivity 
steep. Gaster: petiole think and erect, with erect hairs; gaster reddish-yellow, with thick layer of 
appressed hairs and scattered erect to suberect hairs throughout. 

Queen. — Unknown. 

Male. — Unknown. 

Etymology. — The species epithet, bakwele, is in honor of the Bakwele pygmies who assist- 
ed BLF during his fieldwork in Gabon. 

Measurements.— (Holotype worker) TL: 3.24; HL: 0.902; HW: 0.870; SL: 0.751; ML: 
1.069; GL: 1.272; CI: 96.45; SI: 86.32. 

Discussion. — This new Acropyga species is not only the largest species presently known 
from Africa, but it is also one of the largest in the world. Only four other species are known to 
exceed 3 mm in total length (all are Old World): A. aciitiventris, A. butteli, and A. myops all have 
been observed to exceed 3 mm in total length, whereas A. rubescens has been observed over 5 mm 
in total length. 

The most remarkable attribute of A. bakwele is the presence of a median ocellus. In the exten- 
sive review oi Acropyga by LaPoUa (2004), ocelli were never observed on workers (they are pres- 
ent in queens and males). Unfortunately, with only a single specimen of A. bakwele available for 
study it is impossible to know if the presence of a median ocellus is typical for this species or if the 
specimen is simply an aberrant worker. Nonetheless, its presence is intriguing and the collection of 
a nest series will hopefully clarify the point. 

Nothing is known about the natural history of this Acropyga, except that was collected from 
sifted leaf litter (a method that commonly collects Acropyga) in moist, sandy soil rainforest, near 
an extensive network of marshland. Where the natural history is known, Acropyga are found to nest 
close to soil in leaf litter, rotting logs, and under stones. They form large colonies with thousands 
of workers, and some species are possibly polygynous. 

The relationship of A. bakwele to other species is uncertain, but superficially all African 
species and A. paleartica (known only from Greece) appear closely related. Pending the discovery 
of A. bakwele males, the species remains unplaced in a species-group. The holotype worker resem- 
bles A. arnolcU in many respects. However, A. bakwele is significantly larger than A. aruoldi, pos- 
sesses a longer mandibular apical tooth, and has erect hairs scattered throughout the mesosomal 
dorsum. One interesting characteristic of A. bakwele is that, like A. arnoldi and A. paleartica, it 
possesses a 5:4 palpal formula, a characteristic that may be associated with more basal Acropyga. 
In fact, LaPolla (2004) hypothesized that A. arnoldi represented that most basal extant species. All 
African Acropyga have worker morphologies that suggest a more basal placement. The mandible 
in all species can possess over six teeth, with A. arnoldi known to possess up to nine teeth. A. sil- 
vestrii is known to possess up to seven teeth, although some specimens have been recorded with 
as few as four. 



LAPOLLA AND FISHER: NEW SPECIES OF ACROPYGA FROM GABON 605 



Key to Afrotropical Acropyga species (workers) 

The follow ing key is modified from LaPolla (2004) 

1. Head w idth < 0.55 mm A. silvestrii 

Head width > 0.55 mm 2 

2. Head width < 0.7 mm: total length < 3 mm; erect hairs concentrated on the posterior prono- 

tum: median ocellus absent: southern Africa A. anioldi 

Head width > 0.7 mm; total length > 3 mm: erect hairs scattered throughout dorsum; median 
ocellus present; West Africa A. bah\'ele, sp. nov. 

Acknowledgments 

Funding for the BLF's field research was provided by WWF-US. Travel for JSL to California 
was provided by the Smithsonian Institution's Office of Sponsored Research and National Science 
Foundation under Grant No. INT 9998672 to ELF. We thank April Nobile for creating the images. 
We thank Wojciech Pulowski and Donat Agosti for their reviews of the paper. 

References 

Bolton. B. 1994. Identification guide to the ant genera of the world. Harvard University Press, Cambridge, 

Massachusetts. USA. 222 pp. 
BCnzli. GH. 1935. Untersuchungen iiber coccidophile Ameisen aus den Kaffeefelden von Surinam. 

Mitteilungen der Schweizerischen Entomologischen Gesellshaft 16:455-593. 
Goodman. S.M. R. Hutterer, and P. R. Ngnegueu. 2001. A report on the community of shrews (Mammalia: 

Soricidae) occurring in the Minkebe Forest, northeastern Gabon. Mammalian Biology 66:22-34. 
Johnson, C, D. Agosti, J. H. C. Delabie, K. Dumpert, D. J. Williams, M. von Tschirnhaus, and U. 

Maschwttz. 2001. Acropyga and Azteca ants (Hymenoptera: Formicidae) with scale insects (Stemo- 

rhyncha: Coccoidea): 20 million years of intimate symbiosis. American Museum Novitates 3335:1-18 
La.Polla. J.S. 2004. Acropyga (Hymenoptera: Formicidae) of the World. Contributions of the American 

Entomological Institute 33(3): 1-130. 
L.aPolla, J.S., S.P. Cover, and U.G Mueller. 2002. Natural history of the mealybug-tending ani Acropyga 

epedana, with descriptions of the male and queen castes. Transactions of the American Entomological 

Society 128(3):367-376. 
Weber, N.A. 1944. The Neotropical coccid-tending ants of the genus Acropyga Roger. Annals of the 

American Entomological Society 37:89-122. 



Copyright © 2005 by the California Academy of Sciences 
San Francisco, California, U.S.A. 



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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 

Volume 56. No. 31. pp. 607-617. 5 figs. December 30, 2005 

New and Reconsidered Mexican Acanthaceae XI: 
Justicia in the Yucatan Peninsula 

Thomas F. Daniel^, German CarnevaH^, and Jose L. Tapia Muiioz^ 

' Depmnnent of Botany, California Academy of Sciences, 875 Howard Street, 

San Francisco, California 94103, U.S.A.; Email: tdaniel@calacademy.org. 

- Herbario CICY, Unidad de Recursos Natiirales, Centro de Investigacion 

Cientffica de Yucatan A.C., Calle 43. No. 130. Col. Chiiburnd de Hidalgo 

97200 Merida, Yucatan, Mexico 

Two new species (Justicia edgarcabrerae and J. liizmariae) and a new combination (J. 
leucothamna based on Jacobinia leiicothamna Standi.) are proposed for the acantha- 
ceous flora of the Yucatan Peninsula. Distribution maps, images of pollen, and illus- 
trations/photos are presented for all three species. Studies of Acanthaceae in the 
three states (Campeche, Quintana Roo, and Yucatan) composing the Mexican por- 
tion of the Yucatan Peninsula reveal the presence of at least 38 native species of 
Acanthaceae there. Five of the 13 species of Justicia there are endemic to these states. 

RESUMEN 

Dos especies nuevas {Justicia edgarcabrerae y J. liizmariae) y una combinacion nueva 
ij. leucothamna basado en Jacobinia leucothamna Standi.) se proponen para la flora 
de acantaceas de la Peninsula de Yucatan. Se presentan mapas de las distribuciones, 
imagenes de polen, e ilustraciones/fotos para cada especie. Estudios de las 
Acanthaceae en los tres estados (Campeche, Quintana Roo y Yucatan) que compren- 
den la porcion mexicana de la Peninsula de Yucatan revelan la presencia por lo 
menos de 38 especies nativas de Acanthaceae alii. Cinco de las 13 especies de Justicia 
que crecen alli son endemicos a estos estados. 

Leonard (1936) treated 59 native species in the plant family Acanthaceae from the Yucatan 
Peninsula of southern Mexico and northern Central America. Twenty-seven of these were reported 
from the three states (Campeche, Quintana Roo, and Yucatan) that compose the Mexican portion 
of the peninsula. Recent studies (Daniel, unpublished and this study) reveal the presence of 38 
native species of Acanthaceae in these three Mexican states. Nine of them (24 percent) are endem- 
ic there. Although the acanthaceous tlora of the Mexican portion of the Yucatan Peninsula is not 
especially rich in species, the level of endemism there at that taxonomic rank is significantly 
greater than that noted for several other regions of Mexico, including: Chiapas with 13 percent 
(Daniel 2005aj. "El Bajio" with 5 percent (Daniel and Acosta 2003), Sonora with 3 percent (Daniel 
2004), and the Tehuacan-Cuicatlan Valley with 14 percent (Daniel 1999). However, it is nearly 
equal to the 26 percent endemism reported by Daniel (1997) for species of Acanthaceae in the 
peninsula of Baja California, another very dry region that is not rich in species. The level of 
endemism for the Acanthaceae in the Mexican portion of the Yucatan Peninsula is also high com- 
pared to the estimated 8.2 percent endemism for the total vascular flora of this region (Carnevali et 
al. 2003). 

607 



608 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

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Justicia is the largest genus of Acanthaceae with more than 700 species recognized worldwide. 
It is also the largest genus of Acanthaceae in the Mexican portion of the Yucatan Peninsula with at 
least 13 species native there. Two of these species from the Yucatan Peninsula are newly described 
below and a combination is made in Justicia for the species previously known as Jacobinia leu- 
cothamna Standi. Five species oi Justicia (J. cobensis Lundell. J. clendropila TF. Daniel, J. edgar- 
cabrerae, J. leucothamna. and J. lundeUii Leonard) are endemic to one or more of the three states 
composing the Mexican portion of the Yucatan Peninsula. Another one. J. luzmariae, is known only 
from this region and adjacent northern Belize. 

Ongoing studies toward a comprehensive taxonomic account of the Acanthaceae of the 
Mexican portion of the Yucatan Peninsula (Daniel, in progress), including field and herbarium 
research since 2002, have identified undescribed species (Daniel 2003) and new distribution 
records (Carnevali et al. 2005; Daniel 2005b) for the family. Additional discoveries and a taxonom- 
ic renovation are provided herein. 

Justicia luzmariae T.F. Daniel, Carnevali, and Tapia, sp. nov. 

Fig. 1. 

Type. — Mexico: Quintana Roo: Mpio. Lazaro Cardenas, along hwy. between Kantunilkin and 
Chiquila, 7 km S of Chiquila, 21°22.7'N, 87°22.3'W, 10 m, disturbed evergreen seasonal forest, 25 February 
2003, T. Daniel, G. Carnevali, & J.L. Tapia Miiiloz 10315 (holotype: MEXU!; isotypes: BR!, CAS!, CICY!, 
CIQR!. ENCB!. F!. GH!. K!, MICH!. MO!, NY!, TEX!. UCAM!, US!). 

Frutices usque ad 5 m longi vel alti. Folia petiolata. laminae (ovato-ellipticae vel) ellipticae vel 
subcirculares, 21-90 mm longae, 12-63 mm latae, 1.0-2.6-plo longiores quam latiores. 
Inflorescentia floribus in spicis vel paniculis spicarum. Bracteae obovatae vel obovato-ellipticae, 
3-9 (-14) mm longae, 1-5 (-7) mm latae. Calyx 5-lobus, 6-11 mm longus, lobis homomorphis. 
Corolla viridi-alba vel viridi-lutea et intus maculata, 12-23 mm longa. extus pubescens trichomat- 
ibus eglandulosis. Stamina thecis 1.4-2 mm longis. impariter insertis, pubescentibus, basi cal- 
caratis; pollinis granae 3-aperturatae. Capsula 8.5-14 mm longa. glabra. 

Clambering (sometimes appearing vinelike) shrubs to 5 m long or tall. Young stems sub- 
quadrate to quadrate, bifariously pubescent with retrorse eglandular trichomes 0.1-0.4 mm long. 
Leaves petiolate. petioles to 25 mm long, blades subcoriaceous. somewhat discolorous (lighter 
green abaxially than adaxially), (ovate-elliptic to) elliptic to broadly elliptic to subcircular, 21-90 
mm long. 12-63 mm wide. 1.0-2.6 times longer than wide, rounded to acute to subcordate and 
often asymmetric at base, rounded to acute at apex, surfaces and margin glabrous (or with a few 
eglandular trichomes along midvein on adaxial surface), margin entire, sometimes ± revolute. 
Inflorescence of axillary and/or terminal sessile or pedunculate dichasiate spikes or panicles of 
dichasiate spikes to 132 mm long (including peduncle, if present), axillary spikes (or panicles of 
spikes) (alternate to) opposite. 1 per axil, fertile portion of spikes 7-14 mm in diameter near mid- 
point (excluding flowers), peduncles of spikes to 47 mm long, pubescent like young stems, rachis 
bifariously pubescent with flexuose to retrorse to antrorse eglandular trichomes 0.2-0.5 mm long, 
inflorescence bracts (i.e.. when panicles of spikes present) subulate to elliptic, 2-5 mm long, 1-2 
mm wide: dichasia opposite. 1 per axil. 1 -flowered, sessile. Bracts obdeltate to obovate to obovate- 
elliptic. 3-9 (-14) mm long. 1-5 (-7) mm wide, apically (rounded to) truncate to emarginate. abax- 
ial surface sparsely pubescent with antrorse to antrorsely appressed eglandular trichomes 0.1-0.3 
mm long (trichomes mostly or entirely restricted to midvein). margin ciliate with flexuose to 
antrorse eglandular trichomes. Bracteoles linear to Hnear-elliptic to lunate to lanceolate (sometimes 



DANIEL. CARNEVALI. AND TAPIA: JUSTICIA IN THE YUCATAN PENINSULA 



609 




Figure 1. Justicia Iiizmariae. a. Habit (Crane 509), xO.5. b. Inflorescence (Gomez-Pompa 1352). x3. c. Distal portion 
of stamen witii anther (Gomez-Pompa 1352), xl3. d. Distal portion of style with stigma {Gomez-Pompa 1352), x23. e. 
Capsule (Crane 509). x5, opening capsule (top) and inner side of a single valve (bottom). Drawn by Meg Stalcup. 



asymmetricj, 2.5-7 mm long, 1-2 mm wide, abaxial surface pubescent like bracts. Flowers sessile. 
Calyx 5-lobed, 6-11 mm long, lobes homomorphic, lanceolate, 5-10 mm long, 1-2 mm wide, 
abaxially glabrous or with a few trichomes like those on bracts. Corolla greenish externally, green- 
ish white to greenish yellow internally and with maroon markings on both lips (or with the lower 
lip sometimes light to dark maroon with yellowish green markings), 12-23 mm long, externally 
pubescent with erect to flexuose eglandular trichomes 0.2-0.5 mm long, tube ± abruptly expanded 



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in proximal V3 to ± gradually 
expanded distally, 5.5-10 mm 
long, 3.5-5 mm in diameter near 
midpoint, internally densely 
pubescent near base of stamens, 
upper lip 6-12 mm long, 2-lobed at 
apex, lobes to 0.5 mm long, lower 
lip 6-14 mm long, lobes rounded, 
1-3 mm long, 1.3-2 mm wide. 
Stamens inserted between base and 
midpoint of corolla tube. 8-17 mm 
long, filaments glabrous, thecae 
greenish turning maroon, parallel 
to subparallel. 1.4-2 mm long 
(including basal appendage), equal 
to subequal, unequally inserted 
(overlapping by 0.5-1.2 mm), both 
dorsally pubescent with flexuose 
eglandular trichomes, both with 
blunt basal appendages 0.3-0.7 
mm long (appendage of lower 
theca larger than that of upper 
theca); pollen (Fig. 2) 3-aperturate, 
apertures flanked on each side by 1 
row of insulae, exine reticulate. 
Style 7-19 mm long, proximally 
pubescent with eglandular tri- 
chomes. becoming glabrous distally. stigma 0.1-0.2 mm long, asymmetric, lobes sometimes incon- 
spicuous. Capsule 8.5-14 mm long, glabrous, stipe 2.5^ mm long, head ellipsoid to obovoid, 6-10 
mm long. Seeds 4, plano-convex, 3.2-3.5 mm long, 2.2-2.3 mm wide, surfaces smooth 
(micropapillate). lacking trichomes. 

Phenology. — Flowering: January-March; fruiting: February-April. 

Distribution and habitat. — Yucatan Peninsula of Mexico (Campeche, Quintana Roo) and 
northern Belize (Corozal): plants occur in evergreen seasonal forests ("selva mediana subperenni- 
folia") and tropical subdeciduous forests ("selva baja subcaducifolia"') at elevations of 10-301 m. 

Paratypes. — Mexico: Campeche: Mpio. Calakmul, 3 km E del poblado La Lucha, 18°26'N, 
89°25'W, D. Alvarez & C. Jimenez J. 4205 (CAS); Mpio. Calakmul, 3 km E del poblado Chichonal, carretera 
Xpujil-Escarcega, 18°31'N, 89°32'W, D. Alvarez & C. Jimenez J. 4238 (CAS); Mpio. Calakmul, 4.2 km N 
del poblado La Nueva Vida. 18°50'N, 89°22'W. D. Alvarez & C. Jimenez J. 4369 (CAS); Mpio. Calakmul, 
Puente Papagayo. 25 km N de Xpujil, 18°44'N. 89°24'W, J. Calonico S. et al. 21795 (CAS); Mpio. 
Hopelchen. S de Xpujil rumbo a la frontera, 18°09.5'N. 89°27.5'W. C. Chan 4572 (CICY, GH, MO, UCAM); 
30 km de Sohlaguna. A. Gomez-Pompa 1352 (CAS, CICY). Quintana Roo: Mpio. Carrillo Puerto, 6-10 km 
NE de Felipe Carrillo Puerto, camino a Vigia Chico, E. Cabrera et al. 16373 (CAS): 7-8 km S de Chiquila, 
a lo largo de la carretera Chiquila-Kantunilkin. ca. 21°22'42"N, 87°22'18"W. G Camevali et al. 6733 (CAS, 
CICY. HUH. MEXU. MO. NY UCAM. UJAT. US, XAL); Mpio. Felipe Carrillo Puerto, ca. 6 km NE of 
Felipe Camllo Puerto on road to Vigia Chico. 19°35.9'N, 88°00.3'W. T. Daniel 10298 (BR, CAS. CICY, 
CIQR. K. MEXU, MO, NY, US): Mpio. Lazaro Cardenas, 6 km ENE of San Angel along road (departs 
Kantunilkin-Chiquila hwy. 30 km S of Chiquila) to E. Zapata, 2ri4.2'N, 87°23.2'W, T. Daniel et al. 10316 



Figure 2. Scanning electron micrographs of pollen. a,b. Justiciu 
edgarcabrerae (Cabrera & Diirdii 624). apertural view (a) and interapertur- 
al view (b). c.d. Jiisticia leitcothamna {Leal & Rico-Gray 111), apertural 
view (c) and interapertural view (d). e. Jiisticia hizmariae {Cabrera et al. 
16373). apertural view. f. Jiisticia hizmariae (Gomez-Pompa 1352). interap- 
ertural view. All scales = 2 |im. 



DANIEL, CARNEVALI, AND TAPIA: JUSTICIA IN THE YUCATAN PENINSULA 



611 



(BR. CAS, CICY, K, MEXU). Belize: Corozal: Cerros Maya Ruins, Lowrey's Bight, C. Crane 509 (BRIT, 
LL). 

Vegetatively. Justicia hizmariae appears superficially similar to (and has occasionally been 
identified as) Bravaisia berlandieriana (Nees) T.F. Daniel. Putative relatives of this species are not 
ob\ ious among known species of Justicia from Mexico and Central America, nor does it conform 
to any of the sections of the genus recognized by Graham (1988). Among other species of Justicia 
occurring in the Yucatan Peninsula. J. hizmariae resembles J. lundellii in the shared characters of 
densel) bracteate spike-like inflorescences with prominent bracts, equally 5-lobed calyces, dorsal- 
ly pubescent thecae. and 3-apei1urate pollen. In the latter species, however, the calyx is 2.5-3 mm 
long, the corolla is 7-9 mm long, the pollen is pseudocolpate (lacking insulae), the capsule is 
pubescent, and the seeds are bacculate. 

The species appears to be widespread in eastern and southern regions of Yucatan Peninsula 
(Fig. 3): it has yet to be collected 
in the state of Yucatan. Within J. 
hizmariae. plants from northern 
Quintana Roo (Camevah et al. 
6733. Daniel et al. 10315. and 
Daniel et al. 10316: Fig. 4) differ 
from those from central Quintana 
Roo. southern Campeche. and 
Belize (all other collections 
cited: Fig. 1) by their longer 
corollas (17-23 mm vs. 12-14 
mm), stamens (14-17 mm vs. 
8-9 mm), and styles (17-19 mm 
vs. 7-12 mm). Variation in col- 
oration of corollas is also evident 
in the two populations from 
northern Quintana Roo. There, 
the internal surface of the lower 

lip varies from greenish yellow with maroon markings to light or dark maroon with greenish yel- 
low markings (Fig. 4). The difference in floral length suggests that plants have different pollinators 
in the two regions. In all other features, plants from northern Quintana Roo appear identical to 
those from southern Quintana Roo and Campeche. 

The epithet of this species honors Dra. Luz Maria Calvo Irabien, community ecologist at the 
Centro de Investigacion Cientifica de Yucatan, whose studies and photographs of plants from near 
Kantunilkin led us to this species. 







^^-»-^ 


-^^ 


A 


J- edgarcabrerae 




^^— v"" 


'^ 


H^ 


■ J. leucothamna 


/"^ 




1 


) 


• J. luzmariae 


/ 


/ 




/ 


^ 


/ L.. 

\ 


y 
,y 
y 

\ y 

c 


5 

7 


V 


Tabasco ^- ^- -X 


■ 


•\ Til 


/ 


80 km 


i^w" 


'- , Chiapas ) 1 


Guatemala 


1 Belize / 

i > 



Figure 3. Map of the Mexican portion of the Yucatan Peninsula (with 
states, clockwise from left: Campeche. Yucatan, and Quintana Roo), showing 
distributions of Justicia edgarcabrerae, J. leucothamna, and /. luzmariae. 



Justicia edgarcabrerae T.F. Daniel, Carnevali, and Tapia, sp. nov. 

Fig. 5 

Type. — Mexico: Quintana Roo: brecha a Santa Cruz, 1 km S de Pedro A. Santos, 9 Dec 1980, E. 
Cabrera & G. Durcin 624 (holotype: CAS!; isotype: MEXU!). 

Herbae perennes usque ad 1 m altae. Folia petiolata, laminae ovatae, 13-44 mm longae, 
6.5-21 mm latae, 1.5-2.3-plo longiores quam latiores. Spicae axillares. Bracteae spathulatae vel 
late ellipticae vel subcirculares vel subdeltatae, (5-) 6-9 mm longae, (1-) 2-6.5 mm latae. Calyx 
5-lobus, 3.5-5 mm longus, lobis homomorphis. Corolla luteola, 8.3-11.3 mm longa, extus pubes- 



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Figure 4. Photographs of Jusricia luzmariae (a. b) and J. leiicothamna (c). a. Canievali et al. 6733, xl.8. b. Daniel 
et al. 10316 (maroon form), xl.3. c. Tapia & Chdzaro 1453. x2.5. 



DANIEL. CARNEVALI. AND TAPIA: JUSTICIA IN THE YUCATAN PENINSULA 



613 




Figure 5. Justicia edgarcabrerae. a. Habit {Cabrera & Durdn 624), xO.4. b. Inflorescence nodes (Cabrera & Durdn 
624). x4.8. c. Dichasium (Cabrera 16968 and Cabrera & Durdn 624), x5.8. d. Distal portion of stamen with anther 
{Cabrera & Durdn 624). xl7.3. e. Capsule (Cabrera & Durdn 624). x7.2. f. Seed (Salazar C. 26). x22.6. Drawn by Nadia 
Strasser. 



cens trichomatibus glandulosis et eglandulosis. Stamina thecis 1.1-1.4 mm longis, impariter inser- 
tis. theca supera pubescens trichomatibus eglandulosis, theca inferna basi calcarata; pollinis granae 
3-aperturatae. Capsula 5.5-6.5 mm longa, pubescens trichomatibus eglandulosis. 

Perennial herbs to 1 m tall. Young stems subquadrate, pubescent with erect to flexuose eglan- 
dular trichomes 0.5-1.2 mm long, trichomes disposed throughout but ± concentrated in 2 lines. 
Leaves petiolate, petioles to 8 mm long, blades ovate, 13-44 mm long, 6.5-21 mm wide, 1.5-2.3 
times longer than wide, (rounded to) acute at apex, acute to subattenuate at base, surfaces pubes- 
cent with erect to flexuose to antrorse eglandular trichomes, margin entire. Inflorescence of axil- 
lary (and terminal) pedunculate dichasiate spikes to 53 mm long (including peduncles and exclud- 
ing flowersj, 10-11 mm in diameter near midspike, spikes opposite at nodes, 1-2 per axil, borne 
on peduncles to 5 mm long, rachis ± evenly pubescent with erect to flexuose to antrorse eglandu- 
lar trichomes 0.3-0.8 mm long; dichasia alternate, sessile, 1 -flowered. Bracts opposite to suboppo- 
site, spatulate to broadly-elliptic or subcircular or subdeltate and stalked at base, (5-) 6-9 mm long, 
f 1-) 2-6.5 mm wide, fertile bracts somewhat larger than to conspicuously larger than sterile bracts 



6 1 4 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

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(i.e., bracts subheteromorphic to heteromorphic), rounded to acute at apex, abaxial surface pubes- 
cent with erect to flexuose eglandular and glandular trichomes 0.2-0.5 mm long, margin ciliate 
with trichomes like those of abaxial surface and with eglandular trichomes to 1.3 mm long as well. 
Bracteoles oblanceolate (often asymmetric) to linear, 4-7 mm long, 0.2-1.4 mm wide, pubescent 
like bracts. Flowers sessile. Calyx 5-lobed, 3.5-5 mm long, lobes equal, 2.5^.5 mm long, 0.7-0.9 
mm wide, abaxially and marginally pubescent with erect to flexuose eglandular trichomes 0.5-1 
mm long. Corolla yellowish, 8.3-11.3 mm long, externally pubescent with erect to flexuose eglan- 
dular and glandular (sparse) trichomes 0.1-0.5 mm long, tube 4.2-5.5 mm long (not or scarcely 
expanded distally), 1.5-2.3 mm in diameter near midpoint, upper lip 4-5.3 mm long, apically 2- 
lobed, lobes 0.3-0.5 mm long, lower lip 4.5-6.5 mm long, lobes 1.2-2.5 mm long, 0.8-2.5 mm 
wide. Stamens 4.5-5 mm long, inserted near apex of corolla tube, thecae maroon, 1.1-1.4 mm long 
(including basal appendage), parallel, unequally inserted (overlapping by 0.2-0.3 mm), dorsally 
pubescent with eglandular trichomes, upper theca with a ± inconspicuous basal appendage to 0.2 
mm long, lower theca with a blunt basal appendage 0.5-0.9 mm long; pollen (Fig. 2) 3-aperturate, 
apertures flanked on each side by 1 row of insulae or insulae absent and grains 6-pseudocolpate, 
exine reticulate. Style 7.5-8.5 mm long, proximally pubescent with eglandular trichomes, stigma 
0.2 mm long, lobes not evident. Capsule 5.5-6.5 mm long, pubescent with erect to retrorse eglan- 
dular trichomes 0.1-0.4 mm long, stipe 1.9-2.5 mm long, head ellipsoid with slight medial con- 
striction. Seeds (immature?) 4, plano-convex, 1.1 mm long, 1 mm wide, surfaces tuberculate. 

Phenology. — Flowering: November-January; fruiting: November-January. 

Distribution and habitat. — Yucatan Peninsula of Mexico (Quintana Roo; Fig. 3); plants 
occur in evergreen seasonal forests ("selva mediana subperennifolia") at elevations from near sea 
level tolO m. 

Paratypes. — Mexico: Quintana Roo: Mpio. Felipe Carrillo Puerto, 19 km NW [NE] de F. Carrillo 
Puerto sobre el camino a Vigia Chico, E. Cabrera 16968 (CIQR); Mpio. Felipe Carrillo Puerto, KM 20 carr. 
antigua de F. Camllo Puerto a Vigia Chico, Salazar C. 26 (CIQR). 

The three known collections of this species each note a different color for the corolla (yellow 
for the type, blue for Cabrera 16968, and white for Salazar C. 26). It is possible that each charac- 
terization is at least partially correct, and like several other species of Jiisticia in the region, the 
corollas are cream to yellowish with bluish or purplish markings. 

Among species of Justicia, J. edgarcabrerae appears related to a suite of heteromorphically 
bracteate American species that includes J. chol T.F. Daniel, J. costaricana Leonard, J. nevlingii 
Wassh. & T.F. Daniel, and /. iixpaiiapensis T.F. Daniel (Daniel 2002; Wasshausen and Daniel 
1995). Among these species pollen varies from 2-aperturate (e.g., J. iixpanapensis) to 3-aperturate 
(e.g., /. chol) to 4-aperturate (e.g., J. nevlingii). Justicia edgarcabrerae is especially similar to J. 
chol which occurs in wetter forests of southern Mexico and has corollas that are white to cream- 
yellow with maroon mai'kings (Daniel 1995). It differs from that species by the characters noted in 
the following couplet: 

la. Young stems quadrate to quadrate-sulcate; cauline trichomes with maroon septa; leaves with 
petioles to 35 mm long, blades acuminate at apex; calyx 2.5-3.5 mm long; corolla tube 5.5-7 
mm long; stamens 3.5-4.5 mm long, thecae superposed (separated by a gap up to 0.5 mm 
long): rainforests of Chiapas and Tabasco /. chol 

lb. Young stems subquadrate; cauline trichomes without maroon septa: leaves with petioles to 8 
mm long, blades (rounded to) acute at apex: calyx 3.5-5 mm long; corolla tube 4.2-5.5 mm 
long: stamens 4.5-5 mm long, thecae unequally inserted (overlapping by 0.2-0.3 mm); ever- 
green seasonal forests of Quintana Roo J. edgarcabrerae 



DANIEL. CARNEVALI. AND TAPIA: JUSTICIA IN THE YUCATAN PENINSULA 615 



The epithet of this species is based on the name of the well known Mexican plant collector, 
Edgar Cabrera (see biographical information in Sousa S. and Cabrera C. 1983), whose fine speci- 
mens have enriched knowledge of the Yucatecan flora. Because of the existence of J. cabrerae 
Leonard, named for a different collector, we use both given and family names in this epithet. 

Justicia leiicothamua (Standi.) T.F. Daniel, Carnevali, and Tapia, comb, no v. 

Jacobinia leucothamna Standi.. Field Mus. Nat. Hist., Bot. Sen 8: 44. 1930. 

Type. — Mexico: Yucatan: Silam [= Dzilam Gonzalez, see below], G. Gaumer 1242 (holotype: F!). 

Jacobinia Nees is usually included within Justicia (see Graham 1988), and a combination in 
the latter genus has not previously been made for this species. Justicia leucothamna is apparently 
known only by the six collections from the Yucatan Peninsula listed herein. Thus, it appears to be 
endemic to the Mexican portion of the Yucatan Peninsula (Fig. 3). The affinities of this species 
were not addressed by Standley in the protologue or by Leonard (1936) in a treatment of 
Acanthaceae of the Yucatan Peninsula. In many features (e.g., axillary, secund, and dichasiate 
spikes: four calyx lobes of equal length; whitish corollas: and contiguous but unequally inserted 
and dorsally pubescent thecae. the lower with a prominent basal appendage) the species resembles 
J. sahiiflora H.B.K. of Graham's (1988) section Sarotheca (Nees) Benth. These species differ by 
the distinctions noted in the following couplet: 

la. Leaves to 85 mm long, to 43 mm wide, and 1.2-2.7 times longer than wide, apically acute- to 
rounded- to truncate-apiculate: calyx 3-5 mm long: corolla white with maroon markings, 9-12 
mm long: capsule 8-12 mm long, glabrous J. leucothamna 

lb. Leaves to 170 mm long, to 77 mm wide, and 1.5-4.4 times longer than wide, apically acute to 
acuminate to subfalcate: calyx 5.5-12 mm long: corolla greenish yellow tinged with pink and 
with maroon markings, 12-21 mm long: capsule 14—20 mm long, pubescent. . . J. sahiiflora 

Both of these species have 2-aperturate pollen with trema regions flanked on each side by one 
row of peninsulae or insulae (Fig. 2). 

Martinez S. et al. 3086 J occurs well to the south of other known collections of this species 
(Fig. 3). It was collected in a moister habitat ("selva mediana subcaducifolia") than the collections 
from northern Yucatan ("selva baja caducifolia"), and its stems and leaves are not as densely pubes- 
cent as in plants from the drier regions. On the basis of recent collections, Standley's (1930) 
description of 7. leucothamna can be augmented as follows: corollas white with maroon markings 
on the lower lip (Fig. 4), 9-12 mm long: stamens 4-6.5 mm long, thecae maroon, 1-1.3 mm long; 
capsules 8-11.5 mm long, glabrous; seeds 4, 1.8-2.2 mm long, surface and margin densely tuber- 
culate with conical tubercles. 

The type locality of this species was cited by Gaumer as "Silam." Among his collections of 
Acanthaceae, Gaumer distinguished between "Silam" and "Port Silam." In addition, Millspaugh. 
who worked with Gaumer's collections, distinguished "Silam" from "the port of Silam" 
(Millspaugh 1896); and on his map of the Peninsula (Millspaugh 1896), "Silam" is shown interior 
to the coast. This certainly suggests that "Silam" refers to what appears on modem maps as Dzilam 
Gonzalez, and that "the port of Silam" would refer to what appears on modern maps as Dzilam de 
Bravo (which is situated on the coast, ca. 15 km NE of Dzilam Gonzalez). Thus the type locality 
would appear to be Dzilam Gonzalez. 

Additional specimens examined. — Mexico: Campeche: Mpio. Calakmul, 45 km NW de 
Conhuas, camino a Champoton, 18'^49'N, 90°00'W, E. Martinez S. et al. 30861 (CAS, MEXU). Yucatan: 



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Silam, G. Gaumer2280 (F, GH, MO); Mpio. Rio Lagartos, pasando el Rancho Paraiso rumbo a Las Coloradas, 
2r35'N, 88°10'W, J. Leal & I. Espejel 223 (CICY); Mpio. San Felipe, 16 km despues de Panaba rumbo a 
San Felipe, 21°26'N, 88°15'W, /. Leal & V. Rico-Gray 111 (CICY); Mpio. Dzemul, km 6 de la carretera 
Dzemul-Xtampu, 4 km S del entronque a ruinas de Xtampii, 2ri6.5'N, 89°18.5'W, J.L. Tapia M. & M. 
Chdzaro 1453 (CAS, CICY). 

Acknowledgments 

Daniel's field and herbarium studies were funded by a Franklin Grant from the American 
Philosophical Society in 2003; this award is gratefully acknowledged. We thank Meg Stalcup and 
Nadia Strasser for their fine botanical illustrations; Scott Serata for assistance with the scanning 
electron microscope; Lilia Can, Francisco Chi-May, Celso Gutierrez, Silvia Hernandez Aguilar, 
Gerald Islebe, and Filogonio May for their valuable assistance in the Yucatan Peninsula; and the 
curators of the following herbaria for access to their collections: BRIT, CAS, CICY, CIQR, F, GH, 
LL, MEXU, MO, UCAM. We also thank the reviewers for their thoughtful comments. 

Literature Cited 

Carnevali, G, I. Rami'rez-Morillo, and J. a. GonzalezTturbe. 2003. Flora y vegetacion de la Peninsula de 

Yucatan. Pages 53-68 in P. Colunga Garcia-Marin and A. Larque Saavedra, eds., Naturaleza y sociedad 

del Area Maya: pasado, presente yfiituro. Centro de Investigacion Cientifica de Yucatan, Merida, Mexico. 
Carnevali F, G, J.L. Tapia M., I.M. Ramirez M., R. Duno de Stefano, S. Hernandez A., TF. Daniel, F. 

CoE, J.J.J. Ortiz, N. Diego, L. Can I., and F. May P. 2005. Notes on the flora of the Yucatan Peninsula 

III: new records and miscellaneous notes for the peninsular flora II. Han'ard Papers in Botany 9:257-296. 
Daniel, TF 1995. Acanthaceae. Pages 1-158 in D.E. Breedlove, ed.. Flora of Chiapas, Pt. 4. California 

Academy of Sciences, San Francisco, California, USA. 
Daniel, TF. 1997. The Acanthaceae of California and the peninsula of Baja California. Proceedings of the 

California Academy of Sciences, ser. 4, 49:309-403. 
Daniel, TF 1999. Acanthaceae. In P. Davila A. et al.. eds. Flora del Valle de Tehiiaccin-Cuicatldn, Fas. 23. 

Institute de Biologia, UNAM, Cd. Mexico. 
Daniel, TF 2002. New and reconsidered Mexican Acanthaceae IX. Jitsticia. Proceedings of the Ccdifornia 

Academy of Sciences, ser. 4, 53:37^9. 
Daniel, TF 2003. A reconsideration of Megalostoma (Acanthaceae), a new species, and recognition of a new 

section of Justicia. Proceedings of the California Academy of Science, ser. 4, 54:371-380. 
Daniel, TF. 2004. Acanthaceae of Sonora: taxonomy and phytogeography. Proceedings of the Ccdifornia 

Academy of Sciences, ser. 4, 55:690-805. 
Daniel, TF 2005a. Catalog of Honduran Acanthaceae with taxonomic and phytogeographic notes. 

Contributions from the University of Michigan Herbarium 24:51-108. 
Daniel, TF. 2005b ("2004"). Further range extensions of Mexican Acanthaceae. Polibotdnica 18:1-12. 
Daniel, TF, and S. Acosta C. 2003. Acanthaceae. Pages 1-173 in J. Rzedowski and G Calderon de 

Rzedowski, eds.. Flora del Bajio, Fas. 117. Institute de Ecologia, Centro Regional del Bajio, Patzcuaro, 

Michoacan, Mexico. 
Graham, V.A.W. 1988. Delimitation and infra-generic classification of Justicia (Acanthaceae). Kew Bulletin 

43:551-624. 
Leonard, E.C. 1936. The Acanthaceae of the Yucatan Peninsula. Publications of the Carnegie Institution of 

Washington 46i:l93-23S. 
Millsraugh. C.F 1896. Contribution II to the coastal and plain flora of Yucatan. Field Museum of Natural 

Histoiy. Botanical Series 1:277-340. 
SouSA S., M. and E. Cabrera C. 1983. Listados floristicos de Mexico II. Flora de Quintana Roo. Institute de 

Biologia, UNAM. Cd. Mexico. 
Standley. PC. 1930. Studies of American plants-Ill. Field Museum of Natural Histon: Botcmical Series 

8:3-73. 



DANIEL. CARNEVALI. AND TAPIA: JUSTICIA IN THE YUCATAN PENINSULA 617 



Wassrausen, D.C, and T.F. Daniel. 1995. Justicia nevlingii (Acanthaceae), a new species from Mexico. 
A^mw; 5:114-117. 



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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 

Volume 56. No. 32, pp. 619-623, 4 figs. December 30, 2005 



Vanderhorstia bella, a New Goby from Fiji 
(Teleostei: Gobiidae) 

David W. Greenfield^ and Kenneth R. Longenecker^ 

^ Research Associate, California Academy of Sciences, San Francisco, California and Emeritus 

Professor. University of Hawaii: Email: greenfie@hawaii.edu: Mailing address: 944 Egan Avenue, 

Pacific Grove. California 93950: - Bernice P. Bishop Museum, Honolulu, Hawaii, 96817 

A single individual of a new goby species in the genus Vanderhorstia was collected 
from a fine sand bottom at Vanua Balavu Island, Bay of Islands, in the Northern Lau 
Group of Fiji. The species differs from all other described species in the genus except 
V. mertensi by having 17 dorsal and 18 anal-fin rays. It differs from V. mertensi by 
having about 77 versus 52-62 longitudinal scales and lacking a row of black spots the 
length of its midside. Vanderhorstia bella has much lavender coloration on the head 
and body and many bright yellow spots. 

While conducting a survey of the fishes of Fiji, we collected a single individual of a spectac- 
ularly colored new goby in the genus Vanderhorstia. The specimen was collected using rotenone 
from a fine, silty, sand bottom at a depth of 8.3 m at Vanua Balavu Island, Bay of Islands, in the 
Northern Lau Group of Fiji. 

The genus Vanderhorstia is represented by 12 described species, which are considered to be 
\alid (Winterbottom et al. 2005). 

Materials and Methods 

All counts and measurements follow Winterbottom et al. (2005). Measurements were made to 
the nearest 0.1 mm using dial calipers and are expressed as percentage of standard length (SL). 
Dorsal pterygiophore formula and some other counts were taken from a radiograph. Format gener- 
ally follows Winterbottom et al. (2005) for ease of comparison, and when characters are the same, 
their description is used. The holotype is deposited at the California Academy of Sciences (CAS). 

Species Description 

Vanderhorstia bella Greenfield and Longenecker, sp. nov. 

Figs. 1-^. 

Material Examined. — Holotype: CAS 222208, 70.9 mm SL, Fiji, Northern Lau Group, Vanua 
Balavu Island, Bay of Islands, 17°10.692'S, 179°00.887'W, fine, silty, sand with small coral patch, 8.3 m, 7 
Januar>' 2003, field number G03-22, collected by D.W. Greenfield, K.R. Longenecker, and R.C. Langston. 

Diagnosis. — A species in the genus Vanderhorstia with 17 segmented dorsal-fin rays, 18 seg- 
mented anal-fin rays, about 77 longitudinal scales, and a pointed caudal fin, lacking a row of black 
spots on its side, and having yellow spots on a lavender background on its head and anterior part 
of its body. 

619 



620 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

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Description. — Dorsal-fin elements VI-I,17, all rays branched; anal-fin elements 1,18, all 
rays branched; pectoral-fin rays 18, upper and lowermost rays unbranched; pelvic-fin elements 1,5; 
segmented caudal-fin rays 17, 9 dorsal -i- 8 ventral branched rays; dorsal unsegmented (procurrent) 
caudal-fin rays 7; ventral unsegmented (procurrent) caudal-fin rays 7; longitudinal scales about 77; 
transverse scale rows from anal-fin origin anterodorsally to first dorsal-fin base 20; transverse 
scales from anal-fin origin posterodorsally to second dorsal-fin base 17; predorsal scales absent in 
midline; scales beginning above pectoral-fin base extend posteriorly to insertion of first dorsal fin; 
circumpeduncular scales 12; no scales on pectoral-fin base; scales on prepelvic region embedded 
and difficult to count; gill rakers 4 + 16 on outer surface of first arch; vertebrae 10 -i- 16; dorsal 
pterygiophore formula 3 (2,2,1,1); epural 1; anal-fin pterygiophores anterior to first haemal spine 
2; pleural ribs on third to tenth precaudal vertebrae. 

The following measurements are expressed as % SL: head length 25.2; head width 9.4; head 
depth 13.1; snout length 3.7: eye diameter 5.5; interorbital width 2.0; nape width 7.4; jaw length 
11.1; body depth at origin of first dorsal fin 13.1: body depth at origin of anal fin 11.2; body width 
7.3; predorsal length 29.1; prepelvic length 27.5; preanal length 51.7; caudal-peduncle length 11.9; 
caudal-peduncle depth 7.3; length of first dorsal-fin base 19.5; length of second dorsal-fin base 
41.8; pectoral-fin length 28.2; pelvic-fin length 24.0; length of first dorsal-fin spine 18.3; length of 
second dorsal-fin spine 17.5; length of third dorsal-fin spine 16.7; length of fourth dorsal-fin spine 
27.1; length of fifth dorsal-fin spine 22.4: length of sixth dorsal-fin spine 16.4; length of spine of 
second dorsal fin 8.9; length of first segmented ray of second dorsal fin 12.4; length of longest seg- 
mented ray of second dorsal fin (- 4'h) 13.7; length of anal-fin spine 7.9; length of first segmented 
ray of anal fin 10.2: length of longest segmented ray of anal fin (= 5*) 15.1; length of pelvic-fin 
spine 7.1; length of first segmented ray of pelvic fin 11.5; length of fifth segmented ray of pelvic 
fin 21.6; caudal-fin length 37.1. 

Body elongate and compressed. Head slightly compressed, its width 71.7% of depth. Snout 
very short, its length 67% of eye diameter; snout does not protrude beyond upper lip. Eye dorso- 
lateral, moderately large, its diameter 22.0% of head length; interorbital space narrow its width nar- 
rower than pupil diameter and 8.1% of head length. No distinct, deep trough around eyes from 
interorbital to postorbital regions. No cutaneous ridge along dorsal midline of nape. Gape moder- 
ately oblique, forming an angle of about 28° with body axis. Lower jaw projecting anteriorly 
beyond upper jaw; posterior end of jaws reach to slightly behind posterior eye margin; jaw length 
43.9% of head length. 

Anterior nasal opening a short tube, with the posterior edge slightly longer than the anterior 
edge; posterior nasal opening a large pore, located adjacent to eye. Tip of tongue rounded, anteri- 
or portion free from tloor of mouth. Posteroventral margin of lower lip entire, no mental flap on 
chin. Gill opening wide, extending anteriorly to vertical line through posterior margin of pupil of 
eye; gill membranes attach to very narrow isthmus; no distinct free rear margin. No tleshy projec- 
tions on lateral wing of shoulder girdle. No bony projections along posterior margin of preopercle. 

Caudal peduncle moderately slender, its depth 61.3% of caudal-peduncle length. First dorsal 
fin higher than second dorsal fin; first dorsal fin close to. but not connected to second dorsal fin by 
membrane; fourth spine of first dorsal fin longest, 155.4% of second spine length, not filamentous; 
all dorsal spines slender and flexible; fourth segmented ray of second dorsal fin longest. Origin of 
anal fin on vertical base with first segmented ray of second dorsal fin; height of anal fin slightly 
higher than second dorsal fin; anal-fin spine slender and flexible; fifth anal-fin ray longest. Pectoral 
fin nearly lanceolate, reaching posteriorly to vertical line through base of second dorsal fin between 
spine and first segmented ray; upper and lowermost pectoral-tin rays unbranched. the remainder 
branched. Origin of pelvic fin about midway between posterior edge of opercular membrane and 



GREENFIELD AND LONGENECKER: NEW GOBY FROM FIJI 



621 




Figures 1-4 (CAS 222208); Holotype of Vanderhorstia bella. (1) Full lateral view; (2) Closeup of head and anterior 
portion of body: (3) Closeup of anterior body showing distinctive color markings; (4) Closeup of caudal fin. 



dorsal-fin origin; pelvic fins joined medially by well-developed frenum (between spines) and inter- 
radial membrane (between innermost segmented rays); pelvic frenum moderately thin, with 
smooth posterior margin: all segmented pelvic-fin rays branched. 

Head scaleless. including predorsal; scales cycloid on anterior part of body back to about tips 
of pectoral fins, becoming larger and ctenoid with peripheral cteni posteriorly; no scales on pec- 
toral-fm base; scales overlying basal region of caudal fin all ctenoid. 

Teeth in both jaws unicuspid; upper jaw with outer row of spaced, enlarged, curved caniniform 
teeth and an inner row of small similar teeth, teeth near symphysis enlarged and point posteriorly; 
lower jaw with 1-3 enlarged, curved, spaced, caniniform teeth, two irregular rows of smaller teeth 
medially grading into a single row posteriorly, an innermost row of 2-3 much enlarged curved, 
spaced canines at bend of dentary; no teeth on vomer or palatine. 

Cephalic sensory systems: pore pattern as in Vanderhorstia nannai (Winterbottom 2005, Fig. 
3j. All cephalic sensory-papillae rows uniserial, not forming multiple rows; relatively reduced lon- 
gitudinal pattern of sensory papillae rows on cheek; row a short and reduced, with about four sen- 
sor}' papillae; row b very short, extending back from row a to about one-third distance to preoper- 
cle; row d extending back just past end of maxilla. 

Color of fresh specimen: Background color white, overlaid by lavender in many areas. Side of 
head bright, iridescent lavender covered with many small (about one-third pupil diameter) round, 
bright yellow spots extending from eye back onto pectoral-fin base. Snout and jaws white with a 
slight lavender tinge. A black line in fold between premaxilla and maxilla anterior to eye. Pupil 
black, iris silver with tinges of yellow. Lower side of head white, top of head and nape lavender. 
Sides of body with lavender tinge on upper half (less intense than on head), white on lower half; 
upper half covered with irregularly-shaped yellow spots edged in black; lower half with smaller 
yellow spots without black edges; a series of 13 irregular black vertical lines on midside below sec- 



622 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 32 

ond dorsal fin; a series of 10 dark blotches running from middle of first dorsal-fin base to caudal 
peduncle. Pectoral and pelvic fins clear. First dorsal fin light yellow. Second dorsal and anal fins 
light yellow with distal lavender margins. Caudal fin with light yellow rays and lavender mem- 
branes in between. 

Color in alcohol: Background color cream. Top of snout and anterior portion of premaxilla and 
maxilla dusky, posterior part of jaws cream, a distinct black line between premaxilla and maxilla, 
anterio-ventral to eye. Side of head and pectoral-fin base with numerous round light spots; pupil of 
eye black, iris silver with black dorsal margin. Top of head and nape with scattered small brown 
spots, upper half of body with irregular small light spots surrounded with dark pigment; lower half 
of body and breast cream. Caudal, anal and pelvic fins peppered with small dark pigment spots; 
pectoral fins immaculate; first and second dorsal fins peppered with small dark spots and with 
small round light spots. 

Etymology. — From the Latin belliis, an adjective meaning beautiful, referring to the strik- 
ing coloration of the species. 

Comparisons. — Vanderhorstia bella is in the subfamily Gobiinae because it has a single 
anterior pore in the interorbital area, the lower jaw has more than one row of teeth, both the dorsal 
and anal fins are separate from the caudal fin, and the two dorsal fins are separate. The fish keys 
to the genus Vanderhorstia in Larson and Murdy (2001) because of the following features: la. First 
gill slit open; 2b. Body scaled; 8b. No dermal crest anterior to first dorsal fin; lib. No barbels on 
underside of head; 16b. Dorsal-fin spines thin and flexible; 21b and 22b. Preopercle lacking spines; 
23b. Dorsal-fin origin behind pectoral-fin base; 24b. Cheeks without prominent vertical fleshy 
flaps bearing papillae; 25b. Pelvic fins without fleshy frenum folded forward; 30b. Chin without 
mental frenum; 34b. Head without fine fleshy flaps and bumps; 35b. Head pores present; 39b. 
Pelvic fins completely connected by membrane; 43b. Mouth not small and vertical; 44b. Cheeks 
and operculum naked; 48a. Gill opening extending forward to rear margin of eye; 49b. Head papil- 
lae in a longitudinal pattern; 50a-50b. There is one more anal than dorsal-fin ray, which would key 
to Silhouettea, but because of its very short snout, pointed caudal fin. approximately 77 longitudi- 
nal scales, smooth-edged frenum. and rounded tongue it clearly does not fit the diagnosis of Larson 
and Miller (1986). Also, other Vanderhorstia species have more anal than dorsal-fin rays (e.g., V 
mertensi), thus the key is in error and 50b. was chosen; 51b. No iris lappet and tongue not deeply 
bilobed; 52b. Second dorsal fin and anal fin with 1 spine and more than 10 soft rays; 53b. No dis- 
tinct black ocellus in each dorsal fin; 54b. Caudal fin pointed, longer than head, body with spots, 
and no bright white spot on pectoral fins = Vanderhorstia. As pointed out by Shibukawa and Suzuki 
(2004), there are no derived characters supporting monophyly of Vanderhorstia, and it is separat- 
ed from Ctenogobiops only by caudal-fin length and coloration. 

The number of segmented dorsal and anal-fin rays of V. bella is high (D. 17, A. 18) compared 
to all other described species except V. mertensi Klausewitz which has 16 dorsal-fin rays and 17-18 
anal-fin rays. All other described species have 10-14 dorsal-fin rays and 10-14 anal-fin rays. 
Vanderhorstia bella differs from V. mertensi by lacking its distinctive row of black spots that extend 
down the middle of its sides from the opercle to the caudal peduncle, and by having about 77 ver- 
sus 52-62 longitudinal scales. Its high longitudinal scale count also separates it from all other 
species except V. ambanoro (Fourmanoir). It also differs in its distinctive coloration from all 
described and photographs of undescribed species in the literature. 

Because the holotype was collected in a general rotenone station, we do not know if it associ- 
ates with a shrimp or lives in a burrow; however, many other species in the genus Vanderhorstia 
do. The radiograph of the specimen showed that it had one clam and one snail in its stomach, sug- 
gesting that it may feed on items brought up by a shrimp. 



GREENFIELD AND LONGENECKER: NEW GOBY FROM FIJI 623 



Acknowledgments 

We would like to thank R.C. Langston for assistance in collecting specimens, and Captain B. 
Vasconcellos and the crew of the Mokii Mokiia Hine for assistance in the field. We are grateful to 
J. Seeto. G.R. South. R.R. Thaman, and R.W. Tuxton of the University of the South Pacific, Fiji for 
facilitating our collecting in Fiji. We also thank the Fijian Government and local village chiefs for 
permission to collect fishes. Special thanks to the staff at CAS for providing assistance: D. Catania, 
W.N. Eschmeyer, J. Fong, M. Hoang, T. Iwamoto, as well as an anonymous reviewer. This research 
was supported by National Science Foundation grants INT97-29666 and DEBO- 1027545, and Sea 
Grant Project R/FM-6PD. 

Literature Cited 

Larson. H.K. and RJ. Miller. 1986. Two new species of Silhuettea (Gobiidae) from Northern Australia. 
Japanese Journal of Ichthyology 33(2): 110-118. 

Larson, H.K. antd E.G. Murdy. 2001. Gobiidae. Pages 3578-3603 in K.E. Carpenter and V.H. Niem, eds., 
FAO Species Identification Guide for Fishery Purposes. The Living Marine Resources of the Western 
Central Pacific. Volume 6. Bony Fishes, part 4 {Labridae to Latimeriidae), Estuarine Crocodiles, Sea 
Turtles, Sea Snakes and Marine Mammals. FAO, Rome, Italy. 

Shibuk.'XWa. K. and T. Suzukl 2004. Vanderhorstia papilio, a new shrimp-associated goby from the Ryukyu 
Islands, Japan (Perciformes: Gobiidae: Gobiinae). with comments on the limits of the genus. 
Ichthyological Research 51:113-119. 

WiNTERBOTTOM, R., A. IWATA, AND T. KozAWA. 2005. Vanderhorstia nannai, a new species of burrow-associ- 
ated goby from Palau and the Philippines (Pisces: Gobiidae). Aqua, Journal of Ichthyology and Aquatic 
Biology 9 {3):\Q9-\\A. 



Copyright © 2005 by the California Academy of Sciences 
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Fourth Series 

\blume 56, No. 33. pp. 625-637. 6 figs.. 3 tables. December 30, 2005 

Two New Cardinalfishes of the Indo-Pacific Fish 
Genus Zoramia (Apogonidae) 

David W. Greenfield^, Ross C. Langston-, and Jolin E. RandalP 

1 Research Associate. California Academy of Sciences, San Francisco, California and 

Emeritus Professor, University of Hawaii: Mailing address; 944 Egan Ave., Pacific Grove, 

CA 93950: Email: greenfie@hawaii.edu: - Department of Zoology, University of Hawaii, 

Honolulu, Hawaii, 96822: 3 Bernice P. Bishop Museum, Honolulu, Hawaii 96817 

Two new species of Zoramia (formerly a subgenus of Apogon) are described. Zoramia 
flebila, described from Fiji, has blue spots on the sides, blue teardrop-shaped marks 
under the eyes, and two narrow yellow lines on the midside. It also has a small spot 
surrounded by diffuse melanophores on the caudal peduncle, and lacks an opercu- 
lar spot and dark vertical lines above the anal-fin rays. It has scattered 
melanophores on the breast, pelvic fins, and the entire second dorsal fin, and a line 
of dark pigment along the anal-fin base. There are 27-30 gill rakers, usually 28 or 
29. Zoramia fragilis, previously thought to range from the Indian Ocean into the 
Pacific, was shown to consist of two species; Z. fragilis restricted to Mozambique, 
Madagascar, and the Seychelles, and those in the Pacific Ocean a separate species 
here described as Z. viridiventer. These two species are separated by the number of 
gill rakers and dorsal and anal-fin spine length. 

Fraser (1972) divided the apogonid fish genus Apogon Lacepede into ten subgenera, mainly 
on osteological characters. The subgenera Pristiapogon and Zoramia Jordan were revised by Fraser 
and Lachner (1985), who recognized four species within Zoramia Jordan: A. leptacanthus Bleeker, 
the type species, wide-ranging from the east coast of Africa to the Samoa Islands; A. fragilis Smith 
with a disjunct population, one from Mozambique (type locality), Madagascar, and the Seychelles, 
and the other from Indonesia and the Philippines to the Marshall Islands and Samoa Islands; A. 
gilberti (Jordan and Seale) from the Philippines, Sabah, and Indonesia, east to Palau and Yap; and 
A. perlitus. described as a new species from Palau, Papua New Guinea, Molucca Islands, and the 
Philippines. 

Rodman-Bergman (2004) reviewed the generic and subgeneric classification of the 
Apogonidae. Using external morphology, skeletal characters and a detailed study of the cephalic 
lateralis system, she concluded that Apogon is an unnatural taxon: "Every cladogram generated in 
these analyses showed that the subgenera of these two taxa (Apogon and Pterapogon) were more 
closely related to other genera, than they were to one another." Based on her findings we are treat- 
ing the subgenus Zoramia of Fraser (1972) as a genus. 

While conducting a survey of the fishes of Fiji, we collected individuals of a species of 
Zoramia that we did not recognize. The specimens were similar in color to Z. gilberti, but lacked 
the spot on the opercular flap and have distinctive blue teardrop-like marks on the cheek and blue 
spots on the side of the body above the pectoral fin. In checking comparative material of other 
species of Zoramia, we discovered that the eastern population identified as Zoramia fragilis is a 

625 



626 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 33 

distinct species. The purpose of this paper is to describe these two new species of cardinalfishes. 
We present first the diagnosis of Zoramia based primarily on Eraser and Lachner (1985), followed 
by a revised key to the species of the genus and the descriptions of the two new species. 

Materials and Methods 

Data for the holotype are presented first, followed by the range and mean or mode for all spec- 
imens in parentheses. Measurements were made to the nearest 0.1 mm using dial calipers and are 
expressed as percentage of standard length (SL). Methods of making counts and measurements fol- 
low Eraser and Lachner (1985), except for body depth, which was taken vertically from below the 
origin of the dorsal fin (their measurement from the origin of first dorsal spine to the insertion of 
the pelvic spine is slightly oblique). We also added body width (taken just behind the gill opening), 
predorsal, preanal, and prepelvic lengths, lengths of dorsal- and anal-fin bases, and caudal concav- 
ity, the horizontal distance between the tips of the longest and shortest caudal rays. A microscope 
is needed to see scattered melanophores described in the key for Z. flebila. 

The spines and especially the soft rays of the fins of the species of Zoramia are very fragile 
and often found broken. It is unusual to have a specimen with fully intact fins among older lots of 
museum specimens. Longest caudal-fin ray and caudal concavity measurements of the holotypes 
were taken from photographs in the field before fin rays were broken. The third dorsal-fin spine is 
broken in the holotype of Z flebila. Lateral-line scales often are lost. Eye size as percentage stan- 
dard length versus standard length for Z. flebila and Z gilberti was tested with a two-sample T-test 
(Fig. 3). Specimens used in figure two were from both some of the types and also CAS 2223156. 
Except for Z flebila, Z. viridiventer, and Z fi-agilis, gill raker counts in Table 2 are from Eraser and 
Lachner (1985). Measurements for Z viridiventer were taken from nine BPBM specimens. 
Specimens of the new species have been deposited in the Australian Museum, Sydney (AMS); 
Natural History Museum, London (BMNH); Bishop Museum, Honolulu (BPBM); California 
Academy of Sciences, San Francisco (CAS); Field Museum of Natural History, Chicago (FMNH); 
University of Kansas (KU); National Science Museum, Tokyo (NSMT); South African Institute for 
Aquatic Biodiversity, Grahamstown (SAIAB); and the U.S. National Museum of Natural History, 
Washington, D.C. (USNM). 

Genus Zoramia Jordan, 1917 

Zoramia Jordan, 1917: 46 [type species Apo^o/z graeffi Giinther, 1873, by original description (also monotyp- 
ic) = Apogon leptacanthiis Bleaker, 1856]. 

Diagnosis. — Dorsal rays VI-I.9; anal rays II.9; pectoral rays 13-15 (usually 14); pelvic rays 
1,5; scales finely ctenoid; lateral line complete to caudal-fin base, the pored scales 23-24; median 
predorsal scales 6; scales of body not smaller than lateral-line scales; gill rakers 24-32; bran- 
chiostegal rays 7; vertebrae 10 -i- 14; supraneural (predorsal) bones 3; mouth very oblique, the 
lower jaw strongly projecting; supramaxilla absent; posterior end of maxilla with a distinct notch; 
maxilla with a longitudinal ridge ending just before angle of posterior notch; jaws with two rows 
of very small conical teeth anteriorly, narrowing to one row posteriorly; a single row of very small 
teeth on vomer and palatines, none on ectoptery golds; preopercular ridge smooth, the edge finely 
serrate, becoming smooth dorsally on posterior limb; infraorbitial edge smooth; posttemporal 
smooth; body depth moderately deep, 2.1^.0 in standard length (juveniles more slender, in gener- 
al), and strongly compressed, the maximum width 2.5-3.4 in body depth; caudal fin moderately 
forked; no black stripes (though there may be a dark line along dorsal edge of body); digestive tract 
black. 



GREENFIELD. LANGSTON AND RANDALL: NEW CARDINALFISHES 627 



Key to the Species of Zoramia 

la. No black spot on caudal peduncle: second dorsal spine very long and filamentous, 34-66% SL 
(at SL of 23 mm or more), the third and fourth spines also prolonged (east coast of Africa to 
Samoa Islands) Z. leptacantha (Fig. lA) 

lb. A small black spot midposteriorly on caudal peduncle; second dorsal spine not very long and 
filamentous (except adults of Z. flebila and Z gilberti, but spine length less than 36% SL) . 2 

2a. A prominent to diffuse dark spot posteriorly on opercle; total gill rakers 28-32 (Philippines, 

Indonesia, and western Caroline Islands) Z gilberti (Fig. IB) 

2b. No dark spot posteriorly on opercle: total gill rakers 24 — 30 3 

3a. Four to eight short vertical dark lines above anterior half of anal-fin (Philippines, Indonesia, 

Papua New Guinea, and Palau Z peiiita (Fig. IF) 

3b. No vertical dark lines above anal-fin base 4 

4a. Basicaudal black spot surrounded by a dusky to blackish zone: entire second dorsal fin, breast, 
and pelvic fins with scattered melanophores (microscope needed); a line of melanophores run- 
ning along anal-fin base on body; second dorsal-fin spine 21.8-35.2% SL; body depth usual- 
ly greater than 40% SL (Fiji) Z flebila, sp. nov. (Fig. IC, 2) 

4b. Basicaudal black spot not within a dusky to blackish zone on caudal peduncle: posterior half 
of second dorsal fin. breast, and pelvic fins lacking scattered melanophores: no line of 
melanophores running along anal-fin base on body; second dorsal-fin spine 18.9-24.8% SL; 
body depth usually less than 40% SL 5 

5a. Gill rakers 27-30; second dorsal-fin spine 21.5-24.8% SL; second anal-fin spine 15.1-17.9% 
SL (Mozambique, Madagascar, and Seychelles) Z fragilis (Fig. ID) 

5b. Gill rakers 24—27; second dorsal-fin spine 18.9-21.7% SL; second anal-fin spine 13.0-15.9% 
SL (Philippines to Queensland, east to Marshall Islands and Samoa Islands) 
Z viridiventer, sp. nov. (Fig. IE, 4-6) 

Species Descriptions 

Zoramia flebila Greenfield, Langston, and Randall, sp. nov. 

Figs. IC, 2, Tables 1-2. 

Material Examined. — Holotype: CAS 222057, 40.2 mm SL, Fiji, Northern Lau Group. Vanua 
Balavul Island, Bay of Islands, 17°10.692'S, 179°00.887'W, sand with small coral patch, 8.3 m, 7 lanuary 
2003, field number G03-22, collected by D. W. Greenfield, K. R. Longenecker, and R. C. Langston. 
PaR-ATYPES: BPBM 40152, 38.4 mm SL, collected with holotype; USNM 383148, 35.1 mm SL, collected with 
holotype; CAS 222155, 34.6-39.5 mm (3), Fiji, Vanua Leva, north shore, Great Sea Reef, southwest of Kia 
Island, 16'^18.591'S, 179"02.129'E, isolated coral head in fine sand, 10.8-11.5m, 27 March 2002, field num- 
ber G02-109, collected by D. W. Greenfield, K.R. Longenecker, R. C. Langston, and B. K. Mataitini; FMNH 
116455, 43.6 mm, collected with CAS 222155; BM(NH) 2005.4.25.1, 39.3 mm, collected with CAS 222155; 
AMS 1.43576-001, 36.4 mm, collected with CAS 222155; NSMT-P70721, 41.5 mm, collected with CAS 
222155; SAIAB 75633, 36.1 mm, collected with CAS 222155; BPBM 40153, 33.3 mm, collected with CAS 
222155; USNM 383149, 40.4 mm, collected with CAS 222155. Additional Material Examined: Zoramia 
flebila, CAS 222156 (112), CAS 219847 (1 -DNA = 4024), KU 31970 (1-DNA = 4020), all collected with 
paratypes CAS 222155. Zoramia gilberti: Western Caroline Islands, Yap Island, CAS 83496 (50), CAS 28780 
r40j, CAS 28780 (1), Palau, CAS 85911 (4). Zoramia viridiventer. Solomon Islands, CAS 167414 (9). 
Zoramia leptacantha: Yap Island, CAS 84415 (2), Palau, CAS 84399 (2), Fiji, CAS 222157 (39). Zoramia 



628 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 33 



peiiita: Palau, CAS 30740 (5) paratypes, CAS 
30745 (1) paratype. Zoramis fragilis: Madagascar, 
USNM 211839 (9). 

DiGANOSiS. — A species in the genus 
Zoramia with no distinct dark Hne on the dor- 
sum from the first dorsal-fin origin onto the 
caudal peduncle; no dark lines above insertion 
of anal-fin rays; opercular flap lacking a promi- 
nent or diffuse dark spot; caudal spot small, 
surrounded with many diffuse melanophores 
on caudal peduncle; a peppering of 
melanophores on the breast and pelvic fins, and 
all of the second dorsal fin; distinct blue 
teardrop-like marks on cheek; blue spots on 
side above pectoral fin; two narrow yellow 
lines on midside; an iridescent blue line along 
anal-fin base; total developed gill rakers 27-30. 
usually 28 or 29; second dorsal-fin spine 
21.8-35.2 % SL; body depth 39.7^7.2% SL. 

Description. — Dorsal-fin elements VI-I, 
9; anal-fin elements II. 9 last dorsal and anal-fin 
rays branched to base; pectoral-fin rays 13 
(13-14. usually 14), uppermost two and lower 
two or three unbranched; pelvic-fm rays 1,5, all 
branched; principal caudal-fin rays 17, upper 
and lower unbranched; well-developed gill rak- 
ers 21 + 7 (21-23, usually 22 + 6-7. usually 6, 
total 27-30, usually 28 or 29); pored lateral- 
line scales 24; transverse scale rows above lat- 
eral line 2; median predorsal scales 6; circum- 
peduncular scales 12. 

Proportions (as percent SL; also see Table 
1): Body depth 47.2 (39.7^7.2; 44.1); head 
length 39.7 (38.1^1.8;- 40.0); eye length 14.8 
(14.4-15.7; 15.2); snout length 9.8 (6.8-9.8; 
7.9); bony interorbital width 8.8 (8.4-9.7; 9.2); 
upper jaw length 18.9 (17.7-20.3; 18.8); cau- 
dal-peduncle depth 17.1 (15.4-19.0; 16.8); cau- 
dal-peduncle length 23.0 (18.8-28.5; 24.6); 
predorsal-fin length 37.3 (36.6^1.2; 38.9); 
base of first dorsal fin 19.0 (14.9-19.0; 17.4); 

dorsal-spine lengths— first 10.1 (10.1-14.8; 12.4). second 35.2 (21.8-35.2; 27.3), third [broken in 
holotype. not measured] (19.0-23.2; 21.9), fourth 21.0 (16.6-23.6; 19.7), fifth 15.5 (11.5-15.7; 
13.6), sixth 8.1 (6.8-9.2; 8.0); base of second dorsal fm 20.6 (17.4-24.5; 21.6); spine in second dor- 
sal fm 14.9 (14.9-21.1; 18.0): longest dorsal ray 26.6 (26.6-29.2; 28); preanal length 60.1 
(55.4-63.9; 60.6); base of anal fm 23.7 (20.3-24.0; 22.4); anal-spine lengths— first 4.8 (4.3-5.8; 
4.9). second 13.0 (13.0-16.2; 14.9); longest anal ray broken (25- 27.5; 26.5); pectoral-fin length 




FiGURh 1. Presei\ed Zoraniia specimens: lA: Z lepta- 
cantha. CAS 222157; IB: Z. gilberri. CAS 85914; IC: Z 
flebila, CAS 222057 (Holotype); ID: Z fragilis 
USNM211839; IE: Z viridiventer, CAS84689: IF: Z perli- 
ra, CAS30740 (Paratype). 



GREENFIELD. LANGSTON AND RANDALL: NEW CARDINALFISHES 



629 




Figure 2. Holotype of Zommia flebila, CAS 222057. 



28.3 (24.8-3L1; 26.1); prepelvic 
length 31.8 (31.2-41.0; 35.4); 
pelvic-fin spine 15.1 (15.1-20.8; 
18.3); pelvic-fin length 26.1 
(24.6- 30.0; 26.2). 

Body depth 2.1 (2.1-2.5) in 
SL; body compressed, the width 
2.8 (2.8-3.4) in body depth; dor- 
sal profile of head straight; snout 
length 4.0 (4.0-5.9) in head 
length; orbit diameter 2.7 (2.5- 
2.8) in head length; bony interor- 

bital width 4.5 (4.1— 4-. 7) in head length; caudal-peduncle depth 2.3 (2.1-2.7) in head length; cau- 
dal-peduncle length 1.7 (1.4-2.3) in head length. 

Mouth very oblique, forming an angle of about 50° to horizontal axis of head, the lower jaw 
strongly projecting; maxilla extending to below center of eye, the upper-jaw length 2.1 (1.9-2.4) 
in head length; posterior end of maxilla with a distinct angular notch; dentition as in the genus. 
Tongue narrowly triangular with rounded tip, the upper surface with small papillae. Gill rakers well 
developed, the longest on lower limb nearly half orbit diameter in length. Anterior nostril a small, 
short, membranous tube on side of snout, slightly more than half distance from fleshy edge of orbit 
to median anterior point of upper lip; posterior nostril a nan'ow elliptical opening at level of upper 
edge of pupil, its length about one-fourth pupil diameter. 

Suborbital margin smooth, ending below center of eye; preopercular ridge smooth; posterior 
three-fourths of ventral edge of preopercle and ventral one-quarter of posterior edge finely serrate. 

Color of fresh specimen: Top of head and back greenish gray, overlaid with scattered, small 
melanophores. Two parallel, narrow, yellow lines running along midside from opercle to caudal 
peduncle. Area below yellow lines lighter than dorsum, silvery under pectoral fins and on belly, a 
bluish tinge on area above anal fin. Scattered bluish spots above pectoral fin, overlaying yellow 
lines. Caudal peduncle with heavy concentration of melanophores, forming a dark band at caudal- 
fm base. A small black spot at center of band. Area below eye, preopercle and opercle silvery, 
extending back to join silvery belly. Four relatively large teardrop-shaped blue marks under and 
behind eye, with several more spots extending up along opercular margin. Snout dark green, tip of 
lower jaw with reddish tinge. Pupil of eye black, iris silvery with a greenish band running horizon- 
tally across it at pupil. First two dorsal-fin spines with a reddish tinge, remainder of dorsal fin 
greenish yellow. Pelvic fms reddish. Caudal fin clear except for greenish dorsal and ventral mar- 
gins at base. Anal fin clear with a black band along its base and an iridescent blue line next to it on 
the body. Pectoral fins clear. Often coloration that is blue in life may turn a pink color after the fish 
is dead but still fresh, thus the color in Figure 2 looks pink. Another photograph of the two DNA 
specimens shows a blue color. 

Color in alcohol: Head and body straw yellow. Top of head and sides of body, except area 
under and below pectoral fin, covered with tiny, scattered melanophores. Melanophores more con- 
centrated on caudal peduncle, forming a band. A small black spot about half a pupil diameter cen- 
tered on side of band. Area under eye, preopercle and opercle lacking pigment. Snout and lower 
jaw with scattered melanophores. Area between isthmus and insertion of pelvic fins with scattered 
melanophores. Pupil of eye dark, surrounded by silvery iris. First and second dorsal, caudal, and 
pelvic fms covered with scattered melanophores. Anal fin clear except for a row of melanophores 
along its base. Pectoral fins clear. 



630 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 33 





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GREENFIELD. LANGSTON AND RANDALL: NEW CARDINALFISHES 



631 



Table 2. Total gill-raker counts for species of Zoramia. Counts 
are from Eraser and Lachner (1985), except for Z flebila, Z. fragilis, 
and Z. viridiventer. 

24 25 26 27 28 29 30 31 32 



Z flebilia 
Z. fragilis 
Z. gilberti 
Z. leptacantha 
Z. perlita 
Z. viridiveuter 



24 



3 
26 



12 


12 


1 






7 


28 


2 






10 


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19 


7 


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5 


19 


18 


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14.5 - 



Z. flebila 
Z. gilberti 



Etymology. — The specif- 
ic epithet is an adjective from the 
Latin flebilis (tearful), referring 
to the teardrop-shaped marks on 
the cheek. 

Comparisons. — Zoramia 
flebila differs from Z. leptacan- 
tha by lacking the dark line on 
the dorsum from the origin of the 
first dorsal fm onto the caudal 
peduncle. It also has a caudal 
spot that Z. leptacantha lacks. It 
differs from Z perlita by lacking 
the dark lines just above the 
insertion of some of the anal-fin 
rays. It differs from Z. gilberti by 
lacking either a prominent or dif- 
fuse dark spot on the opercular 
flap, and by having a significant- 
ly (T=-4.14, P=0^000, DF=28) 
smaller eye (Fig. 3). It differs 
from Z. fragilis and Z viridiven- 
ter by having diffuse melano- 
phores on the caudal peduncle in 
addition to a small caudal spot. 

by usually having scattered melanophores on the breast, pelvic fins, and posterior part of the sec- 
ond dorsal fin that are lacking in Z. fragilis and Z viridiventer. It also has a line of dark pigment 
along the anal-fin base that is lacking in both species. Whereas Z fragilis and Z. viridiventer usu- 
ally have black tips on the caudal fin, there is no such coloration in Z flebila. The body is deeper 
(39.7^7.2: 44.1 % SL) in Z flebila than in Z viridiventer and Z fragilis (usually less than 40% 
SL). Zoramia flebila differs from all described species by its distinctive coloration. For a compar- 
ison of gill-raker counts, see Table 2. 

Two DNA tissue samples, 4020 and 4024, are deposited at the University of Kansas. The 
voucher specimens for these samples are 4024 = CAS 219847, and 4020 = KU 31970. 



34 



36 



38 



40 



42 



44 



Standard Length (SL) 

Figure 3. Eye diameter as percentage standard length versus standard 
length. Zoramia flebia open triangles, Z. gilberti closed circles. 



Zoramia viridiventer Greenfield, Langston and Randall, sp. nov. 

Figs. IE, 4-6; Tables 2-3. 

Apogon fragilis (non Smith) Burgess and Axelrod, 1975:1442, lower fig. (Madang, Papua New Guinea). 

Apogon gilberti (non Jordan and Seale) Hayashi, 1980:263, fig. 2 (Ishigaki, Okinawa Prefecture). — Hayashi 
and Kishimoto, 1983: 36, fig. 39 (Iriomote Island). 

Apogon fragilis (non Smith; Russell, 1983:49 (One Tree Island, Capricorn Group, southern Great Barrier 
Reef). — Wass, 1984:13 (American Samoa). — Eraser and Lachner, 1985:43, fig. 1 (Indonesia to Samoa 
Islands). — Eichler and Myers, 1997:136, lower fig. (Ryukyu Islands, Marshall Islands and southern Great 
Barrier Reef). — Okamura and Amaoka, 1997:302, lower right fig., 303 (Amami O Shima Islands). — 
Myers, 1999:130, pi. 53, fig. C (Palau and southern Marshall Islands). 

Zoramia fragilis (non Smith) Randall, 2005:215, middle fig. (western Pacific east to Marshall Islands and 
Samoa). 



632 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series. Volume 56, No. 33 

Material examined. — Holotype: BPBM 32507, 39.2 mm, Papua New Guinea, Madang Province, 
lagoon side of Pig Island (Tab Island), coral patch in 17 m, rotenone, J.E. Randall and PL. Colin, 3 November 
1987. Paratypes: CAS 222277, 10: 32.5^0.0 mm, Caroline Islands, Pohnpei, Tokoteihi, inner reefs border- 
ing lagoon west of pass, depth to 4.5 m, rotenone, R.R. Rofen et al., 1 July 1954; CAS 222278, 8: 35.0-38.0 
mm, Vanuatu, Espiritu Santo, Palikulo Bay, isolated coral head surrounded by sand, 0.5^ m, rotenone, R.L. 
Bolin and R. Persson, 7 October 1958: BPBM 8071. 35: 24.5-36.6 mm, Palau, limestone islet southwest of 
Urukthapel, fringing reef. 9 m, rotenone, J.E. Randall and E.S. Helfman. 11 June 1968: BPBM 9699, 4: 
27.0-31.7 mm, Marshall Islands, Majuro Atoll, lagoon, 2 m, quinaldine, J.E. Randall and A.R. Emery, 30 
March 1970; AMS 1.17086-009, 10: 33.5-38 mm, Papua New Guinea, Madang Harbor, Paeowai Island, 
5°11'S, 145°51'E, 9-11 m.B.B.Collette and party, 25 May 1970; BPBM 15627, 11: 31.5-36.2 mm, Solomon 
Islands. Alite Reef (off Malaita). lagoon coral head, 3 m. rotenone. J.E. Randall and G.R. Allen, 25 July 1973; 
BPBM 15684. 25.9 mm. Solomon Islands, Guadalcanal. Honiara Yacht Harbor, patch reef on mud bottom, 14 
m, rotenone, J.E. Randall and B. Goldman. 2 August 1973; AMS 1.18272-002, 45.0 mm, Australia, Great 
Barrier Reef, Capricorn Group, One Tree Island, lagoon, R.H. Kuiter, 20 September 1974; BPBM 19220, 4: 
33.4-41.3 mm, Indonesia, Molucca Islands, Ambon, Ambon Bay, Poka, adjacent to wreck of ship near dock; 
silty bottom with iron wreckage, 15 m. rotenone. J.E. Randall and G.R. Allen, 16 January 1975; AMS 1.20976- 
008. 10: 34.5^0.0 mm, Australia, Great Barrier Reef. Lizard Island, off Mrs. Watson's Beach, 10-11 m, D.F. 
Hoese and H.K. Larson, 24 November 1978; AMS 1.43600-001, 3: 37.2-38.5 mm, BMNH 2005.5.10.1-3, 3: 
39.2^0.0 mm, BPBM 40155, 6: 33.9^2.3 mm, NSMT-P 70846, 3: 37.9-38.5 mm, SAIAB 75547, 3: 
37.1-38.3 mm, all with same data as holotype; BPBM 39077, 2: 26.5-31.3 mm, Papua New Guinea, New 
Britain, Kimbe Bay. reef off Walindi Plantation, drop-off among branches of sponge. 16 m. quinaldine. J.E. 
Randall and J.L. Earle. 21 August 2002. 

Diagnosis. — A species of Zoramia with only the following dark markings: a small black spot 
midposteriorly on caudal peduncle one-half pupil diameter or more in size: a faint broad dusky 
band on side of snout directly before eye: some specimens with a faint dusky line at base of dorsal 
fins; tips of one or both caudal lobes often blackish; second dorsal-fin spine 18.9-21.7% SL; sec- 
ond anal-fin spine 13.0-1 5. 99f SL: gill rakers 24-27 (rarely 27). 

Description. — Dorsal-fin elements VI-I.9: anal-fin elements II.9: last dorsal-fin and anal-fin 
rays branched to base; pectoral-fin rays 14. uppermost and lower two or three unbranched; pelvic- 
fin rays 1,5, all branched: principal caudal-fin rays 17. upper and lower unbranched: lateral-line 
scales to caudal-fin base 24 (plus one smaller pored scale extending onto base of fin); two near- 
equal scales above lateral line to base of first two dorsal-fin spines, followed by a series of large 
scales in a single row below remaining spines and second dorsal fin. these scales overlapping all 
but nan-ow upper part of scales below: scales below lateral line to origin of anal fin 5: predorsal 
scales 6: circumpeduncular scales 12: total gill rakers on first gill arch 6-1-26 (6-7 -i- 24—27), only 
one with 7 rakers on upper limb (raker at angle included in lower count). 

Body depth 2.6 (2.55-3.3) in SL (specimens less than about 34 mm progressively more slen- 
der): body very compressed, the width 2.8 (2.65-2.8) in body depth: head length 2.55 (2.4-2.55) 
in SL: dorsal profile of head straight: snout length 3.9 (3.95-4.2) in head length: orbit diameter 2.65 
(2.6-2.8) in head length; bony interorbital width 4.45 (4.1^.65) in head length: caudal-peduncle 
depth 2.4 (2.5-2.75) in head length; caudal-peduncle length 1.6 (1.55-1.7) in head length. (See also 
Table 3 for additional porportional measurements.) 

Mouth very oblique, forming an angle of about 50"^ to horizontal axis of head, the lower jaw 
strongly projecting; maxilla extending to below center of eye, the upper-jaw length 2.25 (2.2-2.4) 
in head length: posterior end of maxilla with a distinct angular notch: dentition as in the genus. 
Tongue narrowly triangular with rounded tip, the upper surface w ith small papillae. Gill rakers well 
developed, the longest on low er limb nearly half orbit diameter in length. Anterior nostril a small, 
short, membranous tube on side of snout, slightly more than half distance from fleshy edge of orbit 
to median anterior point of upper lip; posterior nostril a narrow elliptical opening at level of upper 



GREENFIELD. LANGSTON AND RANDALL: NEW CARDINALFISHES 



633 



edge of pupil, its length about 
one-fourth pupil diameter. 

Suborbital margin smooth, 
ending below center of eye: pre- 
opercular ridge slightly irregular, 
but without serrae; posterior 
three-fourths of ventral edge of 
preopercle and ventral half of 
posterior edge finely serrate. 

Origin of dorsal fin over 
third to fourth lateral-line scales, 
the predorsal length 2.4 (2.4-2.5) 
in SL; first dorsal-fm spine 3.9 
(3.45-4.05) in head length: sec- 
ond or third dorsal-fin spines 
longest. 1.9 (1.85-2.15) in head 
length; spine of second dorsal fin 
2.3 (2.35-2.5) in head length: 
first dorsal soft ray longest (sec- 
ond ray nearly as long), 1.5 
(1.45-1.5) in head length; first 
anal-fin spine very short, 7.75 
(7.1-8.4) in head length; second 
anal-fin spine 2.8 (2.5-3.0) in 
head length: first anal soft ray 
longest (second ray nearly as 
long). 1.75 (1.45-1.8) in head 
length: caudal fin 3.1 (2.9-3.1) in 
SL: caudal concavity 3.05 
(3.0-3.15) in head length; pec- 
toral fins L5 (L5-L6) in head 
length, the third or fourth rays 
longest; pelvic fins reaching or 
extending slightly beyond anus, 
the first or second soft rays 
longest. 1.75 fl. 65-1. 85) in head 
length. 

Color of holotype in alcohol 
pale yellowish on head and body, 
a little dusky dorsally on nape, 
along base of dorsal fins, and 
dorsally on caudal peduncle; a 
roundish black spot posteriorly 
on caudal peduncle slightly more 
than half pupil diameter in size; 
scattered melanophores on poste- 
rior half of caudal peduncle but 




Figure 4. Holotype of Zoramia viridiveuter, BPBM 32507. 




Figure 5. Underwater photograph of Zoramia viridiventer taken at site 
where holotype was captured. 




Figure 6. Zoramia viridiventer at Karang Elmas Reef, Halmahera. 



634 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 33 



Table 3. Proportional measurements of type specimens of Zoramia viridiventer as percentage of the 
standard length. 



Holotype 



Paratypes 



BPBM 

32507 



BPBM 

39077 



BPBM 

15627 



BPBM 
40155 



BPBM BPBM 
40155 40155 



BPBM 

40155 



BPBM 
40155 



BBPM 
40155 



Standard Length (mm) 


39.2 


31.3 


33.9 


35.6 


37.5 


38.0 


39.8 


40.4 


42.3 


Body depth 


38.6 


35.5 


38.2 


38.4 


37.7 


38.7 


38.3 


39.5 


38.6 


Body width 


13.7 


13.4 


13.6 


13.7 


14.6 


13.5 


14.2 


14.1 


14.1 


Head length 


39.5 


41.2 


41.3 


41.7 


41.0 


41.1 


39.1 


39.2 


40.0 


Snout length 


10.1 


10.2 


10.0 


10.6 


10.3 


9.8 


9.8 


9.9 


9.8 


Orbit diameter 


14.8 


16.0 


15.5 


15.4 


15.0 


14.6 


14.8 


14.5 


14.2 


Interorbital width 


8.9 


9.2 


9.4 


9.0 


9.1 


9.5 


9.3 


9.6 


9.3 


Upper-jaw length 


17.5 


17.5 


18.3 


17.1 


18.6 


18.7 


17.6^ 


17.4 


17.8 


Caudal-peduncle depth 


16.4 


15.7 


15.9 


16.1 


15.9 


15.0 


15.7 


15.8 


15.8 


Caudal-peduncle length 


25.0 


24.4 


24.6 


24.7 


24.5 


25.5 


25.1 


25.1 


24.4 


Predorsal length 


41.5 


40.0 


41.8 


40.7 


41.0 


41.4 


39.9 


40.1 


39.6 


Base of first dorsal fin 


14.5 


15.0 


15.4 


14.2 


15.2 


15.4 


15.1 


14.8 


14.3 


First dorsal spine 


10.2 


11.9 


10.3 


11.2 


10.1 


10.7 


10.5 


broken 


10.6 


Second dorsal spine 


21.0 


21.6 


19.1 


18.9 


19.0 


18.9 


19.1 


19.0 


21.7 


Third dorsal spine 


20.8 


21.3 


21.1 


19.3 


18.7 


18.7 


18.8 


18.7 


21.0 


Fourth dorsal spine 


16.7 


17.2 


20.3 


16.3 


16.0 


16.1 


15.3 


14.1 


12.2 


Fifth dorsal spine 


11.6 


12.5 


12.7 


11.4 


11.6 


12.1 


11.3 


13.8 


10.9 


Sixth dorsal spine 


7.7 


7.4 


broken 


6.7 


7.8 


7.6 


7.7 


7.2 


7.1 


Base of second dorsal fin 


18.6 


18.5 


18.0 


18.1 


18.1 


18.2 


18.1 


18.8 


17.9 


Spine of second dorsal fin 


17.3 


17.4 


17.7 


17.0 


16.5 


17.4 


17.4 


16.9 


16.3 


Longest dorsal ray 


26.8 


27.2 


27.7 


27.5 


27.8 


27.8 


26.9 


26.9 


27.4 


Base of anal fin 


18.4 


19.1 


18.6 


18.6 


18.6 


18.1 


17.7 


18.5 


18.0 


First anal spine 


5.1 


4.9 


5.7 


5.4 


5.3 


5.0 


5.5 


5.2 


4.8 


Second anal spine 


14.2 


15.9 


15.1 


14.4 


14.0 


13.9 


15.1 


13.0 


15.8 


Longest anal ray 


22.9 


27.9 


broken 


28.3 


24.0 


24.8 


24.9 


22.4 


22.3 


Caudal-fin length 


32.3 


34.5 


broken 


33.6 


33.3 


33.0 


33.7 


32.1 


32.5 


Caudal concavity 


13.0 


13.7 


- 


13.2 


13.2 


13.4 


12.7 


12.6 


13.0 


Pectoral-fin length 


26.4 


25.6 


26.7 


26.9 


26.5 


26.6 


25.2 


25.2 


26.2 


Prepelvic length 


38.4 


40.9 


39.3 


38.5 


40.5 


39.9 


40.4 


38.4 


39.0 


Pelvic spine 


15 8 


15.9 


16.9 


15.4 


15.5 


16.6 


15.6 


14.8 


14.6 


Pelvic-fin length 


22.5 


23.9 


24.0 


22.4 


24.3 


23.7 


23.5 


22.2 


23.7 



far less than the density dorsally on body; a faint broad dusky band on side of snout centered slight- 
ly below middle of eye; spines and rays of fins translucent yellowish, only the first two dorsal-fin 
spines and membranes a little dusky; remaining membranes of fins translucent; tips of caudal-fin 
lobes blackish (faint on lower lobe); the black digestive tract is visible as a faint dark area of the 
abdomen, becoming near-black as the intestine nears the anus. 

Color of holotype when fresh as in Figure 4. The body other than the abdomen is translucent, 
making the vertebral column visible, and the small blue spots are apparent on the operculum and 
upper abdomen. 

Figure 5 is from an underwater photo of an individual of this species taken at the collecting 
site of the holotype. The green area posteriorly on the abdomen often covers more of the abdomen, 
as may be seen in other underwater photographs such as those cited in the synonymy above. 

The two fish of Figure 6 were photographed in 50 m at Karang Elmos Reef, Halmahera 
(0°10'r'N. 128°7'E); note the two vertical blue lines on the side above the pectoral fin. 

Underwater photographs of aggregations of this species may show individuals with or without 
black tips on the caudal lobes. More often than not, at least the upper lobe shows a blackish distal 
end. As mentioned above, museum specimens often have abraded fms, especially the caudal, so 
black tips, had they been present, were lost. 



GREENFIELD. LANGSTON AND RANDALL: NEW CARDINALFISHES 635 



Et^tviology. — The specific epithet is a compound adjective from the Latin viridis for green 
and venter for abdomen, in reference to the green coloration usually present on the abdomen in life, 
at least in adults. 

Comparisons. — Zoramia viridiventer differs from Z. leptacantha by lacking the dark line on 
the dorsum from the origin of the first dorsal fin onto the caudal peduncle. It also has a caudal spot 
that Z leptacantha lacks. It differs from Z perlita by lacking the dark lines just above the insertion 
of some of the anal-fin rays. It differs from Z gilberti by lacking either a prominent or diffuse dark 
spot on the opercular flap. It differs from Z.flebila by lacking diffuse melanophores on the caudal 
peduncle in addition to the small caudal spot, and by usually having black tips on the caudal fin. 
Finally, it differs from Z.fragilis by having fewer gill rakers (24-27 verses 27-30), a shorter sec- 
ond dorsal-fin spine (18.9-21.7 verses 21.5-24.8), and a shorter second anal-fin spine (13.0-15.9 
verses 15.1-17.9). 

Remarks. — We became suspicious that the material reported by Eraser and Lachner (1985) 
as Apogon fragilis Smith, 1961 might contain two species when we noticed the broad gap in the 
distribution of the species shown in their Figure 20 between the Seychelles and Sulawesi, and the 
broad range of the gill-raker counts of A. fragilis in their Table 4. Loans of paratypes of A. fragilis 
from Mozambique and specimens from Madagascar identified as A. fragilis by Eraser and Lachner 
provided a nearly complete separation of gill-raker counts from Pacific specimens (Table 2). This 
difference was reinforced by measurements that demonstrate that the second dorsal and anal-fin 
spines are generally longer in Z. viridventer than in A. fragilis, as shown in our key. 

We are not aware of any color photographs taken of A. fragilis when fresh or alive from the 
two localities for the species given by Smith (1961), Pinda, Mozambique and the Seychelles, or 
from the Madagascar locality reported by Eraser and Lachner. Smith included a painting of the 
species by Margaret Smith with his description of the species. It shows a pinkish-gray fish, becom- 
ing pale bluish gray on the abdomen, with a small black basicaudal spot, strong black stripe on the 
side of the snout and tip of lower jaw, a blackish line at base of the dorsal fins and dorsally on the 
caudal peduncle, black tips on the caudal lobes, and an orange line on the anal fin near the base. 

Kuiter (1998:86) identified two underwater photographs from the Maldive Islands as Apogon 
gilberti, but neither is Zoramia gilberti. The single fish in the figure to the left could be Zoramia 
viridiventer, but without a specimen for study, we cannot be sure. The two fish in the figure to the 
right are gray with a broad iridescent blue-green stripe on the body at the level of the upper end of 
the gill opening, a very small black basicaudal spot, black-tipped caudal lobes, and a tiny black tip 
on the first dorsal fin. They appear to represent an undescribed species. 

The distribution of Zoramia viridiventer is largely as given for the Pacific part of Eraser and 
Lachner's Figure 20 fox Apogon fragilis: Philippines, Indonesia, Palau, Yap and Kapingamarangi 
in the Caroline Islands, southern Marshall Islands, northern Kiribati, Papua New Guinea, Great 
Barrier Reef, Solomon Islands, Vanuatu, and Samoa Islands. They reported the third author and 
associates' collections from Palau (in 1968), Marshall Islands, Solomon Islands, and Papua New 
Guinea. Randall et al. (2004) reported Apogon fragilis from Tonga, but the identification as viridi- 
venter is questionable because of slightly higher gill-raker counts in the limited material available. 

Hayashi (1980) plsiced Apogon fragilis in the synonymy of A. gilberti (Jordan and Scale), type 
locality, Negros. He reported A. gilberti from Ishigaki in the southern Ryukyu Islands; his black 
and white figure is not A. gilberti but appears to be A. viridiventer, in which case it would be the 
first record of the species from Japanese waters. We conclude the same for Hayashi and Kishimoto 
(1983j who reported A. gilberti from Iriomote Island in the Ryukyus. 

Russell (1983) was the first to record this species from Australia (as Apogon fragilis). He hst- 
ed two specimens, AMS 1.18267-005 and 1.18271-002, from One Tree Island, Capricorn Group, 



636 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 33 

southern Great Barrier Reef. The former was found at the Australian Museum by Mark A. 
McGrouther and Sally Reader, who reported it as "dried out beyond retrieval." The latter is a 
species of Canthigaster. The number Russell should have given was AMS 1.18272-002, 45 mm SL. 
It is included above as one of the paratypes of Zoramia viridiventer, as is one lot from Lizard Island 
in the northern Great Barrier Reef. 

This species is usually seen in aggregations in lagoons or bays, sheltering among branching 
corals, sponges, etc. Our collections have come from the depth range of 2-17 m, but as noted 
above, the species may be seen at least as deep as 50 m. 

Acknowledgments 

We thank K.R. Longenecker and B.K. Mataitini for assistance in collecting specimens, and 
Captain B. Vasconcellos and the crew of the Moku Mokua Hine for assistance in the field. We are 
grateful to J. Seeto, G.R. South, R.R. Thaman, and R.W. Tuxton of the University of the South 
Pacific, Fiji for facilitating our collecting in Fiji. We also thank the Fijian Government and local 
village chiefs for permission to collect fishes. We thank M. McGrouther and S. Reader of AMS and 
J.T. Williams of USNM for lending specimens. Special thanks to the staff at CAS for providing 
assistance: D. Catania, W.N. Eschmeyer, J. Fong, M. Hoang, and T. Iwamoto. The authors also 
appreciate the helpful comments of the anonymous reviewers. This research was supported by 
National Science Foundation grants INT97-29666 and DEBO- 1027545, and Sea Grant Project 
R/FM-6PD. 

Literature Cited 

Bleeker, p. 1856. Achtste Bijdrage tot de Kennis der ichthyologische Fauna van Temate (1). Natuurkundig 

Tijchschrift voor, Nederlandsch-Indie 12:191-210. 
Burgess, W., and H.R. Axelrod. 1975. Pacific Marine Fishes. Book 6, Fishes of Melanesia. T.F.H. 

Publications, Neptune City. New Jersey, USA. Pp. 1383-1654. 
EiCHLER, D., AND R.F. Myers. 1997. Korallenfische Zentraler Indopazific. Jahr Verlag, Hamburg, Germany. 

489 pp. 
Fraser, T.H. 1972. Comparative osteology of the shallow water cardinal fishes [Perciformes: Apogonidae] 

with reference to the systematics and evolution of the family. Ichthyological Bulletin of the J.L.B. Smith 

Institute of Ichthyology Rhodes University, Grahamstown (34): 1-105 pp. 
Fraser, T.H., AND E.A. Lachner. 1985. A revision of the cardinalfish subgenera Pristiapogon and Zoramia 

(Genus Apogon) of the Indo-Pacific Region (Teleostei: Apogonidae). Smithsonian Contributions to 

Zoo/ogy (4 12): 1-47. 
Gunther, A.C.L.G. 1873. Die fische der Sudsee (I). Journal des Museum Godeffroy 1:1-128. 
Hayashi, M. 1980. First records of three apogonid fishes from Japan. Japanese Journal of Ichthyology 

27(3):261-267. 
Hayashi, M., and H. Kishimoto. 1983. Fish fauna of Iriomote-Island, Ryukyu Islands. III. Apogonidae 

(Apogoninae). Scientific Report of the Yokosuka City Museum (31):15^6. 
Jordan, D.S. 1917. Notes on Glossamia and related genera of cardinalfishes. Copeia (44):46-47. 
KuiTER, R.H. 1998. Photo Guide to Fishes of the Maldives. Atoll Editions, Apollo bay, Victoria, Australia. 257 

pp. 
Myers. R.F. 1999. Micronesian Reef Fishes, ed. 3. Coral Graphics, Guam, vi + 330 pp. 
Okamura, O.. and K. AmaoK-A.. 1997. Sea Fishes of Japan. Yama-kei Publisher, Tokyo, Japan. 

783 pp. [in Japanese] 
Randall, J.E. 2005. Reef and Shore fishes of the South Pacific, New Caledonia to Tahiti and the Pitcaim 

Islands. University of Hawaii Press, Honolulu. Hawaii, USA. 707 pp. 
Randall, I.E., D.G SvirtH, J.T Willliams, M. Kulbicki, G Mou Tham, P. Labrosse, M. Kronen, E. Clua, 



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.^ND B.S. Mann. 2004. Checklist of the shore and epipelagic fishes of Tonga. Atoll Research Bulletin, no. 

502. 35 pp. 
Rodnlak-Bergman, L.M. 2004. The Cephalic Lateralis System of Cardinalfishes (Percifonnes: Apogonidae) 

and its Application to the Taxonomy and Systematics of the Family. Ph.D. dissertation. University of 

Hawaii, Honolulu, Hawaii, USA. 373 pp. 
Russell, B.C. 1983. Annotated Checklist of the Coral Reef Fishes in the Capricorn-Bunker Group, Great 

Barrier Reef Australia. Great Barrier Reef Marine Park Authority, Special Publication Series 1. 184 pp. 
Smith, J.L.B. 1961. Fishes of the family Apogonidae of the Western Indian Ocean and the Red Sea. 

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1-^3. 



Copyright © 2005 by the California Academy of Sciences 
San Francisco, California, U.S.A. 



638 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 

\blume 56. No. 34, pp. 639-655, 5 pis. December 30, 2005 



Scanning Electron Microscope Studies of Some Early 

Miocene Diatoms from the Equatorial Pacific Ocean with 

Descriptions of Two New Species, Actinocyclus jouseae Barron 

and Actinocyclus nigriniae Barron 

John A. Barron 

MS910, U.S. Geological Siin'ey, Menlo Park, CA 94025; Email: jbarron@usgs.gov. 

Scanning electron microscope (SEM) and light microscope (LM) studies are used to 
propose and describe two new species, Actinocyclus jouseae Barron, sp. nov. and 
Actinocyclus nigriniae Barron, sp. nov. from lower Miocene sediments from equato- 
rial Pacific ODP Site 1219. Parallel SEM and LM studies reveal that Thalassiosira 
bukryi Barron should be transferred to Azpeitia and suggest that Actinocyclus bar- 
ronii Radionova is likely to be a variety of A. radionovae Barron 

During the study of the biostratigraphy of diatoms from lower Miocene (24-17 Ma) sediments 
of equatorial Pacific ODP Site 1219 (7°48.019'N, 142°00.940'W; 5063 m water depth) (Barron, in 
press), two species oi Actinocyclus were observed that were not described by either Barron (1983) 
or Radionova (1991). Description of these new taxa and clarification of the taxonomic relationships 
of three other early Miocene diatoms from ODP 1219 warrants detailed study under LM and SEM. 
The purpose of this paper is to detail the valve ultrastructure of these fossil taxa and resolve their 
taxonomic position. 

Methods and Materials 

For the biostratigraphic study of Barron (in press). Cores 4H through 6H {ca. 24.5 to 17.0 Ma) 
of ODP Hole 199-1219A were sampled at 50 cm intervals, with occasional samples taken at 30 cm 
intervals. Approximately 1 g of material was placed in a 250 ml beaker, disaggregated with a wood- 
en stirring rod, and covered with distilled water. Dilute {ca. 3%) hydrochloric acid was then added 
to remove the calcium carbonate. After the reaction ceased, the sample was washed with distilled 
water and centrifuged at 1200 rpm for 4 minutes duration in order to bring the solution to a neutral 
pH. After completion of the washing process, strewn slides were prepared by transferring the sus- 
pended material with a disposable pipette to a 22 x 40 mm coverslip, which was then dried on a 
hot plate and mounted with Naphrax on a 25 x 75 mm glass slide. 

These slides were examined in their entirety under a light microscope (Leitz Ortholux) at mag- 
nification x500, with identifications checked at xl250. The LM photography was completed using 
a Spot Insight v. 4.0 digital camera on a Leica DML microscope. SEM studies were completed on 
selected samples with a Leo 1450VP microscope. 



639 



640 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 34 

Descriptions of New Species 

Actinocyclus jouseae Barron, sp. nov. 

Plate 1, figs. 1, 4, 5; Plate 2, figs. 1, 2. 

NoMENCLATURAL SYNONYM: Actinocyclus challengeri Jouse in Jouse, (ed.), 1977, pi. 57, figs. 10, 24-25, 36, 
nom invalid (no description). This name is a later homonym of A. challengeri O'Meara, 1876. 

Description. — Diameter 33 to 95 \xm. Linear rays of areolae increasing in size from 8-9 in 
10 |im near the center to about 5 in 10 jitm near the margin. Typically each primary (or sub-pri- 
mary) ray is joined by one secondary ray. High (4-5 )im high) mantle near margin covered by dense 
areolae, about 8-10 in 10 )im. Prominent pseudonodule (circular, about 0.7 |j,m in diameter) locat- 
ed at crest of margin. Valve surface undulated, with a raised marginal region, which occupies about 
one third of the valves diameter. This is followed inward by an abrupt depression and a gentle rise 
to the valve's center. Primary radial areolar rays extend from the valve's center to the raised mar- 
ginal region. These are separated by three shorter, secondary areolar rays. 

Comments. — Actinocyclus jouseae resembles the early middle Miocene diatom, A. ingens 
van nodus Baldauf in Baldauf and Barron (1980) in that the valve is undulated with a raised cen- 
ter and it possesses a dense radial, linear pattern of areolae. Whereas the areolae of A. jouseae 
increase in size from 8 to 9 in 10 |im near the center to about 5 in 10 |im near the margin, the are- 
olae of A. ingens var. nodus decrease in size toward the margin (5 areolae in 10 |im near the cen- 
ter to 9 areolae in 10 |im near the margin). This character gives the areolar pattern of A. jouseae a 
finer, denser appearance than that of A. ingens var. nodus. 

Derivation of name. — In honor of Anastasia P. Jouse, diatomist and pioneer diatom stratig- 
rapher. 

Material examined. — Holotype: CAS accession number 625066, CAS slide number 221091, 
ODP 1219A-4H-4. 58-59 cm (Plate 1, figure 1), Deposited at the California Academy of Sciences, San 
Francisco; Paratypes: CAS slide number 221090, ODP 1219A-4H-4, 8-9 cm (Plate 1, figure 4); CAS slide 
number 221092, ODP 1219A-4H-5, 8-9 cm (Plate 1, figure 5). 

Stratigraphic range. — early Miocene (20.0-19.1 Ma) (Barron, in press). 

Actinocyclus nigriniae Barron, sp. nov. 

Plate 1, figs. 2, 3, 6, 7; Plate 2, figs. 3. 4. 

NoMENCLATURAL SYNONYMS: Cestodlscus sp. 6 of Schrader, 1976, pi. 12, fig. 4.; Cestodiscus kugleri sensu 
Fourtanier, 1991, pi. 1, fig. 5. 

Description. — Diameter 15 to 70 lim. Number of rays: three to four rudimentary rays in 
small specimens to 15 in larger specimens. Areolae decrease slightly in size from 8 in 10 |j.m near 
valve center to 11-12 in 10 )im near margin. Steep mantle, densely areolated 11-12 areolae in 10 
|im. Prominent rounded pseudonodule located near crest of submarginal ring. Valves dimorphic, 
convex and concave, larger specimens tend to be flatter; smaller specimens tend to be domed. 
Distinctive "star-like" hyaline rays, which consist of a primary areolar row beginning near the cen- 
ter of the valve, and three to four additional rays of areolae on either side of the primary row, begin- 
ning at regular distances toward the margin. Note: Similar to Cestodiscus praerapax Radionova, 
1991, pi. IV, figs. 1,12; however, it lacks stripes on margin (N. Radionova, 2005, written commun.) 

Comments. — Actinocyclus nigriniae resembles Cestodiscus kugleri Lohman 1974; however, 
its radial hyaline rays are less step-like in appearance and its valves possess a prominent pseudon- 
odule on their raised, submarginal ring. Actinocyclus nigriniae is also similar to Cestodiscus praer- 



BARRON: NEW SPECIES OF MIOCENE DIATOMS 641 



apax Radionova, 1991, pi. IV, figs. 1, 12; however, it lacks stripes on margin (N. Radionova, 2005, 
written commun.) 

Derivation of name. — In honor of Cathy Nigrini, radiolarian biostratigrapher. 

Material examined. — Holotype: CAS accession number 625068, CAS slide number 221093, 
ODP 1219A-5H-6, 110-111 cm (Plate 1, figure 6). Deposited at the California Academy of Sciences, San 
Francisco. IsoTYPES: CAS slide number 221093, ODP 12 19A-5H-6, 110-111 cm (Plate 1, figure 2); CAS slide 
number 221093. ODP 1219A-5H-6, 110-111 cm (Plate 1, figure 3); CAS slide number 221093, ODP 1219A- 
5H-6. 110-111 cm (Plate 1, figure 7). 

Stratigraphic R.ANGE. — early Miocene (22.7-22.3 Ma) (Barron, in press). 

New Combination 

Azpeitia bukryi (Barron) Barron, n. comb. 

Plate 3. figs. 1-5: Plate 4, figs. 1-5. 

Basionym: Thalassiosira bukryi Barron, 1983:511, plate IV, fig. 1. 

Original description. — "Flat, round valve 20 to 60 \xm in diameter. Hexagonal areolae 
(about 7 in 10 |im) arranged in a sublinear to eccentric pattern in the central Y4S of the valve with 
2 to 4 marginal eccentric rows of progressively smaller areolae (9 to 12 in 10 |im). Areolae pattern 
resembles that of Thalassiosira oestrupii (Ostenfeld) Proshkina-Lavrenko. A small hyaline central 
area about 1-2 jim in diameter is often present, especially in larger forms, commonly containing a 
rounded central nodule. Numerous small pores dispersed over the valve face separated by 3 to 4 of 
the larger areolae. Marginal apiculi separated by 7 small areoale. Thin striated margin (1 um in 
width) with 10 radial striae in 10 |im." 

Holotype.— USNM 348710, Plate IV, fig. 1, sample DSDP 77B-28-6, 28-30 cm 

Emended description. — Azpeitia bukryi (Barron) Barron possesses a ring of weakly stalked, 
rimoportulae opening on the valve/mantle interface (Plate 4, figures 1, 3) that are all similar in 
appearance (Plate 4, figure 3). Although no distinct annulus is present (Plate 3, figures 3-5), larg- 
er valves commonly possess a slightly off-center central process (rimoportulae) that is surrounded 
by a hyaline area (compare Plate 3, figures 1-3). When viewed under the SEM (Plate 3, figures 
3-5), the eroded remains of this central rimoportulae shows little or no external projection (Plate 
3, figure 5) in a manner similar to that oi Azpeitia tabularis (see Figure XIV, IB of Fryxell et al., 
1986). Numerous interlocular pores appear on the valve's surface (Plate 3, figures 1-5; Plate 4, fig- 
ure 4) that are assumed to be rimoportulae. The internal openings of these rimoportulae, however, 
are eroded in the specimens examined so far under the SEM and are not diagnostic (Plate 4, figure 
5). Because satellite pores of strutted processes are normally preserved in fossil material even when 
the tubes of strutted processes have been eroded (Hasle, 1985), it is assumed that these processes 
are labiate processes. 

Areola are loculate with external cribra lying slightly below the valve surface (Plate 4, figures 
2, 4), arguing against placement in Thalassiosira. Elongated areolae separate the tube-like cham- 
bers of the marginal rimoportulae (Plate 3, figures 3^; Plate 4, figure 3). This distinctive margin- 
al structure gives the appearance of being striae in LM (Plate 3, figures 1-2). Girdle bands have 
not yet been observed in the fossil material of A. bukryi. 

Comments. — Azpeitia is "characterized by valves with a nearly central labiate process often 
on the edge of an annulus, a ring of labiate processes on the valve mantle, specialized areolar pat- 
terns of the mantle differing from those on the face of the valve, and two or more (usually three) 
hyaline girdle bands including a wide valvocopula" (Fryxell et al. 1986). 



642 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 34 

The regular ring of marginal rimoportulae, the shallow mantle with a valve structure differing 
from that of the valve face, the loculate areolae with external cribra all support transfer of T. bukryi 
to Azpeitia. Similarly, like many species of Azpeitia, T. bukryi seems to have preferred warmer 
waters during its early Oligocene to early Miocene range (Barron et al. 2004). 

The marginal rimoportulae, structure of the shallow mantle and presence of numerous, scat- 
tered rimoportulae on the valve surface closely resemble those of Azpeitia biannulata Sims in 
Mahood et al. (1993), which was described from the lower Oligocene of Prydz Bay, Antarctica. 

Stratigraphic range. — early OUgocene to early Miocene (33.1-17.5 Ma) (Barron et al. 
2004; Barron, in press). 

Comparison of Actinocyclus barronii Radionova and A. radionovae Barron 

Radionova (1985, 1987, 1991) studied early Miocene diatoms from DSDP Sites 63, 65, 66, 
166, 289, 574, 575, providing SEM illustrations of many taxa and describing five new species, 
Actinocyclus barronii, A. mutabilis, A. praellipiticus, Cestodiscus umbonatus, and C. praerapax. 
Although her species A. barronii closely resembles A. radionovae Barron 1983, Radionova (1991) 
stated that it differed from A. radionovae by its having (1) a considerably smaller undulation of the 
valve. (2) the absence of shortened lines of areolae. (3) the presence of hyaline ribs surrounding the 
central hyaline field. During the study of the early Miocene diatoms of ODP 12 19 A, it became 
clear that further taxonomic study was necessary to distinguish the two species of Actinocyclus. 

Actinocyclus barronii Radionova, 1985 

Actinocyclus barronii Radionova. 1985:72. pi. 1. fig. 1; Radionova. 1991:65, pi. V, figs. 2, 4. 

Description (taken from Radionova [1991] because an English translation for that paper was 
available). — "Valve round, sometimes oval, 60-100 )im, slightly concave. Central part of valve (Vs 
of its diameter) occupied by a flat hyaline field. This field has a polyangle as star-shaped and con- 
nects with the rest of the valve by hyaline ridges, which continue in the line of areolae and reach 
to the margin of the valve. Pseudonodule large, without operculum. On the mantle of the valve 
occur 8-10 rimoportulae, which on the external surface are ended by a side aperture, which is a lit- 
tle smaller than the psuedonodule. Mantle is short (low), margin with rough striae." 

Comments. — Extensive examination of Site 1219 material reveals that specimens assignable 
to A. barronii possess shortened lines of areolae (Plate 5, fig. 2) and appear to only differ from the 
considerable variation in the morphology of A. radionovae (Plate 5, figs. 1, 2-6) by the much- 
reduced undulation of their valves. It is not clear what Radionova (1991) means by hyaline ribs sur- 
rounding the central hyaline field (compare Plate 5, figs. 1-2, 5). Given also that the range of spec- 
imens assignable to A. barronii falls completely within the range of A. radionovae, it would appear 
that A. barronii represents a variety of A. radionovae . This hypothesis would have to be confirmed 
by an examination of Radionova's (1985) type material of A. barronii. 

Stratigraphic r.\nge. — early Miocene (19.9-19.1 Ma) (Barron, in press). 

Actinocyclus radionovae Barron, 1983 

Actinocyclus radionovae Barron, 1983:504. pi. III. figs. 1-3; pi. IV, figs. 4-6; Barron, 1985, pi. 1, fig. 2; 
Radionova. 1991:65. pi. V. fig. 1. 

Description (Barron, 1983). — "circular valve with undulating surface 40 to 100 |im in 
diameter. Hyaline central area 10 to 25 |im in diameter with primary and secondary rows begin- 



BARRON: NEW SPECIES OF MIOCENE DIATOMS 643 



ning at different distances from the valve's center, giving a 'star burst' appearance. Submarginal 
area with eroded labiate processes similar to those of Cestodiscus arranged radially every 7 to 10 
}im. Margin 2 )im wide with 9 to 13 striae every 10 )im. Prominent rounded luminate pseudonod- 
ule located near the margin." 

Comments. — In the present study considerable variation has been observed in forms assigned 
to A. radionovae. Both concave valves with hyaline centers (the type concept) and convex valves 
with centers filled by continuation of the areolar rays appear to occur, especially amongst smaller 
(<50 jim diameter) forms (Plate 5, figures 1, 3-6). 

Stratigraphic range. — early Miocene (22.0-19.1 Ma) (Barron, in press). 

Acknowledgments 

Robert Oscarson, USGS, provided invaluable assistance with the SEM. This manuscript ben- 
efited from the comments of Nora Radionova and the critical reviews of Mary McGann and Scott 
Starratt. I am also grateful to Ms. Pat Sims of the British Museum for her very helpful comments 
on the transfer of Thalassiosira bukryi to Azpeitia. An anonymous reviewer also offered helpful 
comments. 

Literature Cited 

Baldalt, J.G, and J. A. Barron. 1980. Actinocyclus ingens var. nodus: a new stratigraphically useful diatom 

of the circum-North Pacific. Micropaleontology 26:103-110. 
Barron, J. A. 1983. Latest Oligocene through early middle Miocene diatom biostratigraphy of the eastern 

tropical Pacific. Marine Micropaleontology 7:487-515. 
Barron, J. A. 1985. Late Eocene to Holocene diatom biostratigraphy of the equatorial Pacific Ocean, Deep 

Sea Drilling Project Leg 85. Initial Reports of the Deep Sea Drilling Project 85:413^56. 
Barron, J. A. (In press.) Diatom biochronology for the early Miocene of the equatorial Pacific. Stratigraphy. 
Barron, J.A., E. Fourtanier, and S.M. Bohaty. 2004. Oligocene and Earliest Miocene Diatom 

Biostratigraphy of Site 1220, ODP Leg 199, Equatorial Pacific. Proceedings of the Ocean Drilling 

Program, Scientific Results 199(204): 1-25. {<http://www-odp.tamu.edu/publications/199_SR/204/ 

204.htm>) 
Fourtanier, E. 1991. Diatom biostratigraphy of equatorial Indian Ocean Site 758, ODP Leg 121. Proceedings 

of the Ocean Drilling Program, Scientific Results 121:189-208. 
Fryxell, GA., pa. Sims, and TP Watkins. 1986. Azpeitia (Bacillariophyceae). Related genera and promor- 

phology. Systematic Botany Monograph 13:1-74. 
Hasle, GR. 1985. The fossil diatom Thalassiosira ornica, n. sp., Micropaleontology 31:280-284. 
Hasle, GR., and pa. Sims. 1986. The diatom genus Coscinodiscus Ehrenb. C. argus Ehrenb. and C. radia- 

tus Ehrenb. Botanica Marina 29:305-318. 
JousE, A. P., editor-in-chief 1977. Atlas of Microorganisms, Bottom Sediments of the Oceans {Diatoms, 

Radiolarians, Silicoflagellates, Coccoliths). Nauka, Moscow, USSR. 196 pp. 
LoHMAN, K.E. 1974. Lower middle Miocene marine diatoms from Trinidad. Verhandlungen der 

Naturforschenden Gesellschaft im Basel 84:326-360. 
Mahood, A.D., J. A. Barron, and PA. Sims. 1993. A study of unusual, well preserved Oligocene diatoms 

from Antarctica. Nova Hedwigia Beihefte 106:243-267. 
O'Meara, E. 1876. Report on the Irish Diatomaceae. Proceedings of the Royal Irish Academy 2:235^25. 
Radionova, E.P 1985. Lower Miocene diatoms of the tropical zone from the western part of the Pacific 

Ocean. Izvestia Geological Sciences 7:62-74. Nauka, Moscow, USSR, [in Russian] 
Radionova, E.P 1987. Diatom morphology of genus Cestodiscus from lower middle Miocene depositions of 

the tropical zone of the Pacific Ocean. Methods of Zonal Stratigraphic Work-up According to 

Microorganisms. Micropaleontology Edition 29:141-154. Nauka, Moscow, USSR, [in Russian] 
Radionova, E.P. 1991. Stratigraphy of Neogene sediments in a tropical area of the Pacific Ocean based on 

diatoms. Academy of Sciences of the USSR, Transactions 456:1-107. Nauka, Moscow, USSR, [in Russian] 



644 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 34 

ScHRADER, H.-J. 1976. Cenozoic planktonic diatom biostratigraphy of the southern Pacific Ocean. Initial 

Reports of the Deep Sea Drilling Project 35:605-671. 
Sims, P.A., G.A. Fryxell, and J.G. Baldauf. 1989. Crucial examination of the diatom genus Azpeitia: Species 

useful as stratigraphic markers for the Oligocene and Miocene Epochs. Micropaleontology 3 5 (4): 293-307. 



Copyright © 2005 b\' the California Academy of Sciences 
San Francisco. California, U.S.A. 



BARRON: NEW SPECIES OF MIOCENE DIATOMS 645 



Plates 



646 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 34 



Plate 1 

1 Actinocyclus jouseae Barron n. sp. Holotype, CAS slide number 221091. pseudonodule at 5 o'clock, ODP 
1219A-4H-4. 58-59 cm. 

2, 3 Actinocyclus nigriniae Barron n. sp.. Isotypes. CAS slide number 221093. pseudonodules at 10 o'clock and 
1 o'clock. ODP 1219A-5H-6. 110-111 cm. 

4a,4b Actinocyclus jouseae Barron n. sp.. Paratype, CAS slide number 221090. low and high focus, pseudonod- 
ule just below 3 o'clock. ODP I219A-4H-4. 8-9 cm. 

5 Actinocyclus jouseae Barron n. sp.. Paratype, CAS slide number 221092, larger form with more complex rays, 
pseudonodule at 9 o'clock, ODP 1219A-4H-5. 8-9 cm. 

6 Actinocyclus nigriniae Barron n. sp.. Holotype. CAS slide number 221093. pseudonodule at 1 o'clock, ODP 
1219A-5H-6, 110-111 cm. 

7 Actinocyclus nigriniae Barron n. sp., Isotype. CAS slide number 221093, pseudonodule at 2 o'clock, ODP 
1219A-5H-6, 110-111 cm. 



BARRON: NEW SPECIES OF MIOCENE DIATOMS 



647 




648 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

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Plate 2 

Scale bars for Figs. 1, 2a, 3a, 4a = 20 |im; for Figs. 2b, 3b, 4b = 5 jim. 

1 Actinocyclus jouseae Barron n. sp., external view of valve, Isotype CAS accession number 625066, ODP 
1219A-4H-4, 58-59 cm. 

2a Actinocyclus jouseae Barron n. sp., internal view of valve, Isotype CAS accession number 625066, ODP 
1219A-4H-4, 58-59 cm. 

2b Close-up of Fig. 2a showing pseudonodule and eroded labiate processes on steep mantle. 

3a Actinocyclus nigriniae Barron n. sp., external view of valve with concave center, Isotype, CAS accession num- 
ber 625068, ODP 1219A-5H-6, 110-111 cm. 

3b Close-up of margin of Fig. 3a showing pseudonodule. 

4a Actinocyclus nigriniae Barron n. sp., internal view of valve, Isotype, CAS accession number 625068, ODP 
1219A-5H-6, 110-111 cm. 

4b Close-up of internal opening of pseudonodule of Fig. 4a. 



BARRON: NEW SPECIES OF MIOCENE DIATOMS 



649 




650 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

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Plate 3 

Azpeitia bukryi (Barron) Barron n. comb., LM and SEM photos. 

1, 2 LM views of valve, ODP 1219A-6H-1, 108-109 cm, scale bar = 20 jam. 

3 SEM. External view of valve ODP 1219A-5H-I, 108-109 cm. 

4 SEM, External view of valve, showing possible eroded central process, ODP 1219A-4H-4, 58-59 cm. 

5 Detail of fig. 4 (enlarged 3X). 



BARRON: NEW SPECIES OF MIOCENE DIATOMS 



651 




652 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

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Plate 4 

Azpeitia bukiyi (Barron) Barron n. comb.. SEM photos. 

1 Internal view of eroded marginal labiate processes. ODP 1219A-5H-1, 108-109 cm. 

2 Detail of margin, ODP 1219A-5H-1, 108-109 cm. 

3 Internal view of eroded valve, ODP 1219A-4H-4, 58-59 cm. 

4 Detail of cribra and small openings, ODP 1219A-5H-1. 108-109 cm. 

5 Detail of Fig. 3 (enlarged 2X). 



BARRON: NEW SPECIES OF MIOCENE DIATOMS 



653 




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Plate 5 

Scale bars for all figs. = 20 |am 

1 Actinocyclits radionovae Barron, external SEM. ODP 1219A-4H-4. 58-59 cm. 

2 Actinocyclus barronii Radionova. external LM. pseudonodule at 1 o'clock. ODP 1219A-4H-4, 58-59 cm. 

3 Actinocyclus radionovae Barron, internal SEM. showing mushroom-shaped labiate processes, and internal 
opening of pseudonodule at 3 o'clock, ODP 1219A-4H-4, 58-59 cm. 

4 Actinocyclus radionovae Barron, Holotype of Barron, 1983, USNM 348702, DSDP 495-33-5, 72-76 cm. 

5 Concave and convex specimens of A. radionovae. s.l. ODP 1219A-4H-4, 58-59 cm. 

f> Actinocyclus radionovae Barron, pseudonodule at 5 o'clock, ODP 1219A-4H-4, 58-59 cm. 



BARRON: NEW SPECIES OF MIOCENE DIATOMS 



655 








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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 

\'olume 56. No. 35. pp. 657-667. 17 figs.. 4 tables. December 30, 2005 



A New Species of Discothyrea Roger from Mauritius and 

a New Species of Proceratium Roger from Madagascar 

(Hymenoptera: Formicidae) 

Brian L. Fisher 

Department of Entomology, California Academy of Sciences, 875 Howard Street, 
San Francisco, California 94103 USA; bfisher@calacademy.org 

The worker of Discothyrea berlita sp. nov. from Mauritius is described. This is the 
first record of the genus from Mauritius. D. berlita is known from a single locality, 
Le Pouce. a small sanctuary of native ants on an island overrun with invasive ant 
species. Proceratium avium is recollected at Le Pouce and is the senior synonym of 
Proceratium avioide de Andrade (syn. nov.). The practice of manually removing alien 
plants from native forest plots in Mauritius is not advised for the Le Pouce forest 
patch because this practice facilitates the establishment of invasive ants, which elim- 
inate native ants. Proceratium google sp. nov. is described from Madagascar. 

Keywords: Conservation, Discothyrea, Formicinae, Hymenoptera, 
Invasive, Madagascar, Mauritius, Proceratium. 

In May 2005. 1 joined a team of Malagasy ant specialists on an expedition to the island of 
Mauritius, where we conducted an ant inventory and a search for indigenous species. The status of 
the remaining native species of Mauritius was called into question by P.S. Ward (1990). In inspired 
literary prose, he described, as W.L. Brown (1974) did earlier, the alarming difficulty of finding 
native species. Habitat destruction and introduced ants and plants dominate the landscape, pushing 
native ants up to and possibly over the brink of extinction. 

Mauritius has had a long history of exploitation, habitat modification and extinction. With the 
extinction of the dodo in 1681, 80 years after humans first arrived on Mauritius, colonizers contin- 
ued to modify habitat at an alarming rate (Lorence and Sussman 1986). The dense Mauritian forests 
were converted into tea and sugar plantations in the 19* century. During this time, habitat modifi- 
cation on Mauritius reached to almost every corner of the island (Safford 1997). Mauritius is an 
instructive example of what could happen to other insular environments, such as Madagascar, if 
habitat destruction is left unchecked. On Mauritius, as on Madagascar, invasive plant and animal 
species pose major problems. Once established, many invasive ants in Mauritius may be virtually 
impossible to eradicate, thus preventing the return of native ants (Holway et al. 2002). 

The known native ant fauna of Mauritius includes 18 valid species, with 9 endemic to the 
island (Table 1). It is interesting that the endemic ants are all confined to upland forest. One could 
conclude that Mauritius has few endemics all of which are on mountaintops. On the other hand, 
these endemics could be the only remaining examples of a much richer endemic fauna that disap- 
peared with the destruction of the lowland forest. The discovery of a new genus record on Le 
Pouce, suggests that there are more species to discover on the island and that Le Pouce is a surpris- 
ing sanctuary of taxonomically peculiar endemic ants. 

657 



Tapinoma pallipes Smith, 1876) are excluded. 
Species in bold are endemic to the island. 



658 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 35 

The site encompasses a rugged and spectacular Table 1 List of valid names for native ants 
mountain chain above the industrial city of Port recorded from the island of Mauritius. Native ants 
Louis in north-west Mauritius. The main ridge runs restricted to Rodrigues {Monomohum elongatum 

approximately east to west, and three long spurs ^"'.^^:.A^l^:„I'^^"L'!;!!/?o^i!:^^ ^."''*:_,.^,!^^' 
extend northward. Major peaks include Pieter Both 
(823 m), Le Pouce (812 m) and Montagne Ory 

(c.700 m). Le Pouce captures moisture from the Camponoms muvsus Roger, IS63 

prevailing wind and clouds, resulting in the pres- Camponoms gmndidieri Ford, 1886 

r ■ 1 I r 1 rx^T • • 1 1 CrewaWPaster sewellii Forel, 1891 

ence oi native cloud-forest there. This is the only r>- ^i u /v 

-' Dicotltiyea berlita sp. nov. 

remaining area of native vegetation, although Hxpoponem johannae (ForA, \mi) 

native plants are scattered throughout the range. Nesomyrmex gibber Dor)i^t\\orgt,\9A6 

Exotic vegetation dominates, most notably a scrub OcheteUus vinsoni (Donisthorpe, 1946) 

r , /r> • 7- 7 • X 1 Plieidole picataForel. 1891 

Of strawberry guava (Psidium cattleianum) and Pheidole tarda F)or(.,^hor^e, 19A1 

privet {Ligustrum robustrum) — but grassland and piagiolepis madecassa Forel, 1892 

Eucalyptus plantations also occur. The best native Pristomyrmex bispinosiis (Donisthorpe. 1949) 

forest' found during our trip, and also the place of Pristomyrmex browni Wang. 2003 

1 ^ J . 11 1 Pristomyrmex trispinosm {F>on\i,ihorr)Q. \9A6) 

greatest number of endemic ants, was a small patch „ ,. . „ im/i 

° ^ Proceratiiim avium Brown, 1974 

of forest, less than one hectare in area, just at the Pseudolasius dodo (Donisthorpe. 1946) 

southeast face of the peak. Based on our survey Soienopsis mameri Dornsihor^t, 1946 

results across the island, this forest patch on Le Strumigenys agetos Fisher, 2000 

„ . , , . . ^ r r ^ Teckuoimnnex ivimrosae F>orn'ii\\orvie, \9A9 

Pouce IS the only remaining forest refuge for these 

mountain endemics of Mauritius and should receive high conservation priority. 

Materials and Methods 

This work is based on ant inventories in Mauritius from 25 May-31 May, 2005. During that 
period, we visited Le Pouce Mt., Pieter Both Mt.. and Calebasses Mt. in the Moka Range, and 
Camizard Mt., and Brise Mt. in the Bambous Range. We also collected at Basin Blanc, He aux 
Aigrettes, Cocotte Mt., and Petite Riviere Noire Mt. Ants were collected using general hand search 
techniques and leaf litter extraction. The work in Madagascar is based on arthropod surveys in 
Madagascar that included over 6,000 leaf litter samples. 4,000 pitfall traps, and 8,000 additional 
hand collecting events throughout Madagascar in 1992 through 2004 (Fisher 2005). The species 
described here was collected as part of an inventory of Reserve Speciale d'Anjanaharibe-Sud 
organized by Steve Goodman (Fisher 1998). 

All species and type material examined in this study have been imaged and are available on 
AntWeb {www.annveb.org). Material was deposited at Cahfornia Academy of Sciences, San 
Francisco (CASC) Museum of Comparative Zoology, Harvard University, Cambridge, 
Massachusetts (MCZC). and British Museum of Natural History (BMNH). 

Digital images (Fig. 1-17) were created using a JVC KY-F75 digital camera and Syncroscopy 
Auto-Montage (v 5.0) software. All metric measurements were taken at 80x power with a Leica 
MZ APO microscope using an orthogonal pair of micrometers and recorded to the nearest 
0.001mm and rounded to two decimal places for presentation. The accuracy of the micrometers 
was tested against a 0.01 mm microscope micrometer before and after measurements. 
Measurement indices and their abbreviations used in the paper are based on those used by Ward 
(1988). Size and the shape of the IV abdominal segment are the most important characters for the 
identification and delimitation of Proceritiinae species. 



FISHER: NEW' FORMICIDAE FROM MAURITIUS AND MADAGASCAR 659 



List of Abbreviations Used 

HL Head length: maximum longitudinal length from the anteriormost portion of the projecting clypeus to 

the midpoint of a line across the back of the head. 
fr\V Head width: maximum width of head, including the eyes, and is taken behind them. 
CI Cephalic index: HW/HL x 100. 

SL Scape length: maximum chord length excluding basal condyle and neck. 
SI Scape index: SL/HW x 100. 
W'L Weber's length: in lateral view of the mesosoma, diagonal length from posteroventral corner of 

mesosoma to the farthest point on anterior face of pronotum, excluding the neck. 
LS4 Length of abdominal sternum IV as described in Ward ( 1988). 
LT4 Length of abdominal tergum IV as defined in Ward ( 1988). 
IGR Index of gastric reflexion: LS4/LT4 

Discothyrea berlita Fisher, sp. nov. 

Fig. 1-4. 

Type ]VL\TERIAL. — Holotype: Worker. Mauritius: Le Pouce Mt., Moka Range, 20°ir55"S, 
057°31'44"E, 750 m, closed vegetation, 25 May 2005 (coll. B.L. Fisher et al.) Collection code: BLF12148, 
specimen code: CASENT0007016 (CASC). 

Type worker measurements: HL 0.57, HW 0.52, CI 91, SL 0.36, SI 70, LS4 0.08, LT4 0.43, 
WL 0.64 IGR 0.19. 

Diagnosis. — The following character combination differentiates berlita from all its con- 
geners: scrobe absent, fused frontal carinae projecting perpendicular to the plane of the clypeus, 
expanding apically. not forming a thin lamellae; propodeal angle without acute teeth or spines; 
anterior margin of petiole concave when viewed from above. 

Et\'MOLOGY. — The specific name is an arbitrary combination, to be treated as a noun in appo- 
sition. 

Worker Description. — Form of head, mandibles, and body as shown in Figures 1-4. 
Antennae 10-segmented; medium segments extremely short and not distinct when viewed with less 
than lOOx magnification; scape expanded apically, reaching mid-point of head. Eyes with 2 or 3 
facets. Without depressed scrobal area. Palpal segmentation requires dissection and thus was not 
determined. Mandible masticatory margin concave, with two teeth, sharp apical tooth and smaller 
acute basal tooth. Propodeal angle without teeth or acute angles; declivitous face of propodeum 
concave. Petiole thick, with lateral margins on anterior face; anterior margin concave when viewed 
from above. Petiole with distinct convex subpetiolar process. Abdominal segment III longer than 
broad. 

Head and mesosoma densely punctulate; petiole sculptured as mesosoma, abdominal segment 
III with sparse punctures; punctures evanescent on abdominal segment IV. Integument generally 
opaque, except shiny for impunctate areas of metasoma. 

Body, including mandible and appendages, covered with dense fine, very short whitish decum- 
bent pubescence, becoming sparse on abdominal segment III, and dense and nearly erect on 
abdominal segment IV. 

Color testaceous red. 

Dlstribution. — The single specimen was collected in a leaf litter sample in the only remain- 
ing patch of dense native vegetation near the summit of Le Pouce. Samples from other nearby 
mountain tops, Pieter Both (823 m), Calebasses (c.600 m), did not uncover any endemic 
Proceritiinae. 

Comments. — The African species of Discothyrea fall into two groups: (1) those with the 



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Figure 1-4. Discothyrea bedita worker: holot\pe CASENT0007016. 



FISHER: NEW FORMICIDAE FROM MAURITIUS AND MADAGASCAR 661 



clypeo-frontal fusion flat topped and broad and with a depressed scrobe region, and (2) those in 
w hich the process forms a simple convex or angular vertical plate and lack a depressed scrobe 
region (Brown 1958). The Discothyrea of Madagascar belong to the first group. D. berlita is most 
similar to those in the second group, but is distinct in that the vertical plate does not form a thin 
lamella, but is expanded apically (Fig. 3). 

Proceratiiim avium Brown, 1974 

Figs. 5-13. 

Proceratiiim avium Brown, 1974: 71, figs. 1 and 2 (worker, gyne and male). Mauritius: Le Pouce Mt, 700- 

800 m. Native forest, 1 Apr. 1969 (coll. W.L. Brown) [examined] AntWeb MCZTYPE32216 (MCZC) [de 

Andrade 2000:75] 
Proceratiiim avioide de Andrade 2003: 78, figs 37, 38 (worker, gyne and male). Mauritius: Le Pouce Mt, 700- 

800 m. Native forest, 30 March 1969 (coll. W.L. Brown) [examined] AntWeb MCZTYPE35017 (MCZC). 

New synonymy [see justification below] 

During the trip to Le Pouce on May 25 and 30, seven new collections of Proceratiiim from Le 
Pouce were recorded (Table 2). Because of the small size of the forest patch, only two complete 
colonies were collected. For the other colonies we encountered, only a few foragers were removed. 
As Brown (1974) observed, foragers were returning to nests with what appeared to be spider eggs. 
In this case, they carried the eggs in the mandible, and did not support the eggs with the recurved 
gaster (Brown 1980). Baroni and de Andrade (2003) suggest the recurved gaster serves a phrag- 
motic function, but I did not observe the recurved gaster being used to plug up the ant nest entrance. 

Table 2. Collection of Proceratiiim avium on 25 and 30 May 2005 at Le Pouce Mt., Moka Range, 
20°11'55"S, 057°3r44"E. 750 m, closed vegetadon. 

Collection Habitat Caste 

BLF12011 foraging on NiLxia venicillata wilh Pristomyniiex bispinosus 1 w 

BLF12014 foraging on Niixia verticillata with Pristomynnex bispinosus 2 w 

BLF12I36 ex rot pocket, Nuxia verticillata, L 5 m above ground 1 erg Q, 127 w 

BLF12137 ex rot pocket, Nuxia verticillata, L 5 m above ground 1 erg. 352w 

BLF12139 foraging on NiLxia veiticillata with Pristomynnex bispinosus 2 w 

BLF12140 foraging on Nuxia verticillata 8 w 

BLF12142 ioraging on Nuxia verticillata 2w 

Of note is the fact that colony (BLF12137) included 352 workers, one ergatoid queen, and no 
males. Based on the colony size data reported in Baroni and de Adrade (2003), this is the largest 
colony size recorded for Proceratium. Collections in May by Brown in 1969 included males. All 
nests encountered were located in Nuxia verticillata Lamark (Loganiaceae), with entrances about 
1 .5-2 m above ground. This tree was also the preferred nesting site for Pristomyremx bispinosus. 
This tree, called bois maigre in Mauritius, has gnarled and twisted trunks. It is endemic to 
Mauritius and Reunion and appears to be the sole nesting site for Pristomyremx bispinosus and 
Proceratium avium. The high winds that are common on Le Pouce abrade the twisted and inter- 
twined trunks and branches. This action damages the tree at the contact point between intersecting 
branches and leads to the creation of a rot pocket and nesting site. 

Three collections of Proceratiimi avium (BLF120I1, 12014, and 12139) were foragers follow- 
ing Pristomyremx bispinosus. These two species are very similar in color and general appearance. 
Brown in 1969 also observed this behavior. It is unclear why Proceratium is interspersed among 
the foraging workers of P. bispinosus. Conservation of either of these species should include fur- 
ther investigation of potential beneficial interactions between the species. 



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HpP^ 


■ 




W^ • 


• '^^^B 




r '-^'^' 'i 


n^^H 




1. t'i 
1 


" 





i 


^mI 


^ 


^^^^Q3 


fe 




Figures 5-13 Profile, head in full-face \ie\\ and mesosoma in dorsal view of Procerariuiu avium workers: Figures 5-7: 
C.ASENT0059014. BLFI2136. collected May 30. 2005; Fig. 8-10: MCZTYPE32216 holotype of Pwcerariiim avium col- 
lected 1 Apr. 1969: Fig. 11-13: MCZTYPE35017 holotype of Proceratium avioide collected 30 March 1969. 



JrSTiFiC.\TlON OF Synonym\\ — Brown (1980) collected three series of Proceratium at Le 
Pouce in 1969. one on March 30. and two on April I. The latter were located less than 500 meters 
from the March 30 collecting site. He described both of these samples as Proceratium avium 
(Brown 1974). De Andrade (Baroni Urbani and de Andrade 2003) reexamined these three collec- 



FISHER: NEW FORMICIDAE FROM MAURITIUS AND MADAGASCAR 663 



tions and determined that they represent two species, P. avioide and P. avium. She based this on the 
observation that P. avium differs from P. avioide by the less impressed sculpture, by the denser 
pilosity. and by longer antennal scapes {P. avium SI 87.3-88.6, P. avioide SI 81.8-83.3). 

The measurements of Brown and de Andrade are not consistent, especially for the P. avioide 
material she examined. Brown noted measurements for the three collections (workers n = 19) as 
HL 0.92-0.98, HW .091-0.98, CI 96-101 SL 0.90-0.99. Brown did not calculate SI. De Andrade 
notes that for her avium: HL 1.05-1.12, HW .090-0.94, CI 84.5-85.7 SL 0.93-0.97, SI 87.3-88.6 
and P. avioide. HL 1.10-1.16, HW .092-0.97, CI 82.1-85.1, SL 0.90-0.96, SI 81.8-83.3. Note that 
CI for Brown ranged from 96-101, while for De Andrade, CI ranged from 82.1-85.7. 

One possible reason for these differences is the differences of HW and SL definitions. Based 
on the definitions presented above, I re-measured the type material using a calibrated micrometer 
(see Methods above). Measurements are pre- x^ble 3. Measurements and scape index of type 

sented in Table 3. These measurements con- material and new collections. MCZTYPE32216 is the 
firm the relative differences between the holotype of Pwceratium avium, MCZTYPE35017 is 
Brown collections. However, when samples the holotype of Proceramwn av/oz<ie. 
from the seven new collections are included, 
these differences become less distinct. The 
seven collections in the study, have even less 
impressed sculpture than P. avium, similar 
pilosity as P. avium, and longer antennal 
scapes then both P. avium and P. avioide (SI 
98-103). Based on this study of Brown's 
material and the new collections in this study, 
I identify all these collections as one species. 

The variation observed in these collec- 
tions is interesting in such a small area. It is 
possible that because P. avium has ergatoid 

queens, and disperses presumably by budding with low dispersal ability, the complex topography 
of Le Pouce contributed to the observed variation. The possible restriction of the remaining popu- 
lation to the single forest patch at the base of the southeast peak, however, could severely limit the 
observed variation in the future. 

Proceratium google Fisher, sp. nov. 

Figs. \^\1. 

Type Material. — Holotype: Worker. Madagascar: Antsiranana, 11.0 km WSW Befingotra, 
Reser\e Speciale Anjanaharibe-Sud, 14°45'S, 049°27'E, 1565 m, 16 Nov 1994 (coll. B.L. Fisher) sifted litter, 
montane rainforest, Collection code: BLF1232(6) — CASENTO 100367, (CASC) Paratypes: 2 workers with 
same data as holotype but with specimen codes CASENT010068 (BMNH), CASENT0100369 (MCZC): 1 
worker 9.2 km WSW Befingotra, Reserve Speciale Anjanaharibe-Sud, 14°45'S, 049°28'E, 1280 m, 5 Nov 
1994 rcoll. B.L. Fisher), CASENTO 100370; (CASC); and 1 worker same as latter but collected at 1200 m on 
9 Nov 1994, CASENTO 100371 (CASC). 

Diagnosis. — The following character combination differentiates P. google from all its con- 
geners: abdominal segment IV tergite evenly rounded posteriorly, without concave impression near 
apex and not hypertrophied; truncate median clypeal lobe; low nodiform petiole without peduncle 
but with blunt anteroventral tooth; fore tibia with a basal spine, frontal carinae separate and diverg- 
ing posteriorly; posterior dorsum of mesosoma and propodeal spines granulate-foveolate. P. google 
is easily distinguished from P. diplopyx, the only other described Proceratium from Madagascar, 



Specimen number 


HW 


SL 


SI 


MCZTYPE35017 


0.97 


0.92 


95 


MCZTYPE32216 


1.01 


0.96 


96 


CASENT0055844 


0.98 


1.01 


103 


CASENT0055842 


0.99 


1.00 


101 


CASENT0059012 


0.97 


0.99 


102 


CASENT00590I3 


1.03 


1.01 


98 


CASENT0059026 


1.00 


1.01 


101 


CASENT0059030 


0.99 


1.01 


102 


CASENT0059029 


1.01 


1.00 


98 


min 


0.97 


0.92 


95 


max 


1.03 


1.01 


103 



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17 



1 mm 



Figures 14-17. Proceraiium google worker: holot\pe C.A.SENTO 100348. 



FISHER: NEW FORMICIDAE EROM MAURITIUS AND MADAGASCAR 665 



Table 4. Worker measurements: maximum and minimum based on all five Proceratium google 
specimens. 



Specimen 






















CASENT# 


HL 


HW 


CI 


SL 


5/ 


WL 


LS4 


LT4 


IGR 


0100367 


Holotype 


1.21 


1.02 


84 


0.80 


79 


1.34 


0.20 


0.85 


0.23 


0100370 


Parat>pe 


1.24 


1.07 


86 


0.92 


86 


1.49 


0.18 


0.79 


0.23 


0100371 


ParaU'pe 


1.24 


1.04 


84 


0.86 


83 


1.46 


0.20 


0.77 


0.26 


0100368 


Paratope 


1.15 


1.03 


89 


0.84 


82 


1.36 


0.19 


0.79 


0.23 


0100369 


Parat}'pe 


1.20 


1.05 


87 


0.83 


79 


1.41 


0.17 


0.79 


0.22 




min 


1.15 


1.03 


84 


0.83 


79 


1.36 


0.17 


0.77 


0.22 




max 


1.24 


1.05 


89 


0.86 


83 


1.46 


0.20 


0.79 


0.26 



b\ the shape of the tergite of the abdominal segment IV. In P. diplopyx, the tergite is with a deep 
conca\e notch near apex. 

Etymology. — Named in recognition of the support from the Google company. I hope that 
Google will continue applying its talent to serve data relevant to the biodiversity community, con- 
servation planners, and the general public. By creating a "Zoogle,"" Google could help achieve free 
and democratic access to taxonomic and biodiversity data on species. P. google is also suspected 
to be a specialist egg predator of spiders, which is also why this ant is aptly named after Google — 
for the ability to hunt down obscure prey. The specific name is an arbitrary combination, to be treat- 
ed as a noun in apposition. 

Worker description. — Form of head, mandibles, and body as shown in Figures 14-17. In 
full-face view, posterior margin of head rounded, not concave; sides of head more or less straight 
medially; in profile, dorsal margin marginate. Mandible with 4 teeth. Palpal formula 4, 3. Antennae 
I2-segmented, scape does not reach posterior margin of head. Median clypeal lobe raised and 
notched medially. Eye a single, large, clear, convex facet that projects beyond the margin of the 
head in full-face view. 

Mesosoma in dorsal view pear-shaped, broader across pronotum than across propodeum. 
Metanotal grove unmarked. Propodeal spines granulate-tuberculate; declivitous face of propodeum 
concave, ending basally with an upturned tooth. Petiole longer than wide; subpetiolar process 
forming an obtuse tooth at midlength. Tibial spur present on each leg. Claws on all legs slender, 
simple. 

Abdominal segment IV tergum evenly rounded posteriorly, without concave impression near 
apex. 

Head, mesosoma. petiole, and abdominal segment III with dense granulate-foveolate sculp- 
ture. In contrast, abdominal segment IV predominantly smooth and shiny but with sparse foveae. 
Declivitous face of propodeum shiny smooth. 

Body covered with abundant pilosity consisting of fine, curved, tapered, yellow-white setae. 

Queen, male and larvae unknown. 

Distribution. — Known only from an isolated mountain in Northeastern Madagascar, 
Reser\'e Speciale Anjanaharibe-Sud, 14°45'S, 049°27'E, collected at an elevation of 1565 m. 
Collections in nearby mountains such as Marojejy did not locate any specimens of this species. 

Conservation 

Arthropods present several challenges to those dedicated to their conservation. First, they are 
small and inconspicuous, and thus often forgotten during the conservation planning process. 
Second, arthropods are overwhelmingly diverse and as a whole, barely known. Is it pragmatic to 



666 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 35 

develop a conservation strategy for a fauna we scarcely know? Third, because arthropods show a 
remarkable level of local endemism, they will require strategies and policies that are different from 
those developed to conserve birds and plants. A case in point is Mauritius. 

Conservation in Mauritius is heavily biased to bird and plant preservation (Safford and Jones, 
1998; Fowler et al 2000. Nicholas et al. 2004). Land management practices are tailored to benefit 
plants and birds, but not invertebrates. They are fighting the battle to protect the dwindling patch- 
es of native vegetation and bird populations. For plants, this includes the establishments of 
Conservation Management Areas where alien plants are manually removed (Dulloo et al 2002). In 
these plots, weedy vegetation is removed up to three times a year. The active removal of large num- 
ber of weedy plants, however, creates large areas of bare soil and understory (Dulloo et al 2002, 
pers. obs.). This disturbance facilitates the establishment of invasive ants, at the expense of any 
remaining native ants (pers. obs.). 

The small, one-hectare patch of native forest left on Le Pouce could be destroyed for native 
ants if an active weeding program is initiated. The closed vegetation is essential for the survival of 
the endemic Discothyrea, Pristomyremx, and Acropya, which thrive in the cold, moist understory. 
With weeding and increased insolation and disturbance, the invasive ants that surround this small 
patch would quickly move in and destroy this ant sanctuary. 

An alternative approach to the manual weeding strategy would be to plant native trees around 
this patch, including Nuxia verticillata, which is home to Proceratium and Pristomyrmex. The goal 
would be to create a dense closed canopy of natives around this patch without disturbing the patch 
itself. Over time, the effective size of this patch could expand. We also advise that future collec- 
tions of endemic ants in Mauritius avoid collecting entire colonies. 

Mauritius has shown that once invasive ants take hold, there is almost no way to return the 
land to native ants and healthy arthropod communities (pers. obs.). Therefore, in Madagascar, land 
managers must monitor for invasive arthropods. Even though remnant patches of forest may be 
preserved, invasion by aggressive exotic ants may drive native ants locally extinct. One of the sim- 
plest and most effective methods is to track the presence or absence of invasive ants. In this 
approach, targeted collecting can be performed in habitats and microenvironments most likely to 
harbor invasives. 

Acknowledgments 

This work was supported by the National Science Foundation under Grant No. DEB-034473 1 
to B.L. Fisher and P.S. Ward. Fieldwork that provided the basis for this work could not have been 
completed without the support of Lorch Lach. Andy Suarez, and Vikash Tatayah from the 
Mauritian Wildlife Foundation. I am also grateful for the Arthropod Inventory Team (Balsama 
Rajemison. Jean-Jacques Rafanomezantsoa, Chrislain Ranaivo, Coco Randriambololona, 
Hanitriniana Rasoazanamavo, Nicole Rasoamanana, Clavier Randrianandrasana, Valerie 
Rakotomalala, and Dimby Raharinjanahary) who helped collect and process this material. I thank 
April Nobile for creating the images. 

References 

Baroni Urbani, C. and M.L. De Andrade. 2003. The ant genus Proceratium in the extant and fossil record 
(Hymenoptera: Formicidae). Museo Regionale di Scienze Natiirali, Monografie 36: 1^80. 

Brown, W.L. 1958. Contributions towards a reclassification of the Formicidae. 2. Tribe Ectatommini. Bulletin 
of the Museum of Comparative Zoology at Hanard College. 118:175-362 



FISHER: NEW EORMICIDAE FROM MAURITIUS AND MADAGASCAR 667 



Brow"N, W.L. 1974. A remai-kable new island isolate in the ant genus Proceratium (Hymenoptera: 

Formicidae). Psyche 81:70-83. 
Brown, W.L. 1980. A remarkable new species of Proceratium, with dietary and other notes on the genus 

(Hymenoptera: Formicidae). Psyche 86:337-346. 
Dltlloo, M.E., S.P. Kell, and C.G. Jones. 2002. Conservation of endemic forest species and the threat of 

invasive species. International Forestry Review 4:277-285. 
Fisher. B.L. 1998. Ant diversity patterns along an elevational gradient in the Reserve Speciale 

d"Anjanaharibe-Sud and on the western Masoala Peninsula, Madagascar. Pages 39-67 in S.M. Goodman, 

ed., A Floral and Faunal Inventory of the Reseire Speciale d'Anjanaharibe-Siid, Madagascar: with 

Reference to Elevational Variation. Fieldiana: Zoology, n.s., 90:1-246 
Fisher. B.L. 2005. A Model for a Global Inventory of Ants: A Case Study in Madagascar. Pages 78-89 in N.G. 

Jablonski and M.T. Ghiselin, eds.. Biodiversity: A Symposium Held on the Occasion of the 150''' 

Anniversai-y of the Califoniia Academy of Sciences June 17-18, 2003. Proceedings of the California 

Academy of Sciences, ser. 4, 56(Suppl. I). 
FowTER. S.V.. S. Ganeshan, J. Mauremootoo, and Y. Mungroo. 2000. Biological Control of Weeds in 

Mauritius: Past Successes Revisited and Present Challenges. Pages 43-50 in Neal R. Spences, ed.. 

Proceedings of the X''' International Symposium on Control of the Weeds, 4-14 July 1999. Montana State 

Uni\ ersity. Bozeman. Montana, USA. 
Holway, D.A., L. Lach, A.V. Suarez, N.D. Tsutsui, and T.J. Case. 2002. The causes and consequences of 

ant invasions. Annual Review of Ecology and Systematics 33:181-233. 
Lorence, D.H., ANT) R. W. Sussman. 1986. Exotic species invasion into Mauritius wet forest remnants. 

Journal of Tropical Ecology 2:147-162. 
Nichols. R.. L. Woolaver. and C. Jones. 2004. Continued decline and conservation needs of the endangered 

Mauritius olive white-eye Zosterops chloronothos. Otyx 38:291-296. 
Safford, R.J. 1997.A survey of the occuiTence of native vegetation remnants on Mauritius in 1993. Biological 

Consenation 80:181-188. 
Safford, R.J.. and C.G. Jones. 1998. Strategies for Land-Bird Conservation on Mauritius. Consen'cition 

Biology 12:169-176. 
Ward, P.S. 1988. Mesic elements in the western Nearctic ant fauna: taxonomic and biological notes on 

Amblyopone, Proceratium, and Smithistruma (Hymenoptera: Formicidae). Journal of the Kansas 

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4:3-5. 



Copyright © 2005 by the California Academy of Sciences 
San Francisco. California, U.S.A. 



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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 

\blume 56. No. 36. pp. 669-674, 3 figs., 1 table. December 30, 2005 



A New Species of Deepwater Snake Eel, 

Ophichthus pulliis (Anguilliformes: Ophichthidae), 

from Angola and Guinea-Bissau 

John E. McCosker 

California Academy of Sciences, 875 Howard Street, San Francisco, California 94103; 
Email: Jmccosker® calacademy. org 

Ophichthus pulliis, a new species of snake eel, subfamily Ophichthinae, is described 
from specimens trawled in deep water (106-154 m) off Angola and Guinea-Bissau. It 
is most similar to Ophichthus aphotistos from Taiwan and O. crueiitifer from the 
western Atlantic but differs from them and other elongate, deepwater species of 
Ophichthus in its vertebral number (149-153), snout condition, pectoral-fin shape 
and size, and dark coloration. 

Keywords: Ophichthidae; Ophichthus pullus sp. nov.; Angola, 
Guinea-Bissau; Ophichthus aphotistos; Ophichthus cruentifer; snake eels. 

Recent deepwater assessment cruises off the eastern tropical Atlantic by the RA^ Dr. Fridtjof 
Nansen have uncovered a new species of snake eel, genus Ophichthus, of the ophichthid subfami- 
ly Ophichthinae (sensu McCosker 1977). The Nansen specimens were made available to a group 
of ichthyologists that met at an FAQ workshop in Tenerife, Canary Islands, during July 2004. The 
specimens had been frozen and thawed, and although somewhat tawdry, they were ultimately rec- 
ognized by the author as an undescribed species. Tomio Iwamoto returned to Angola aboard the 
Nansen in April 2005 and saved two superb specimens of the new species. In order that the name 
of that eel become available for the upcoming publication of the FAO Living Marine Resources of 
the Eastern Central Atlantic (Kent Carpenter, editor), I herein describe the new species and com- 
pare it to the closely related deep-water ophichthines Ophichthus aphotistos and O. cruentifer. 

Most ophichthids occupy habitats shallower than 100 m, ranging from coral reefs to sand and 
mud substrates, entering rivers and estuaries. Recent deep-water trapping, trawling, and sub- 
mersible captures of ophichthids have uncovered a number of new species hving at depths as great 
as 1300 m (McCosker et al. 1989; McCosker 1999; McCosker and Chen 2000). Although most 
ophichthids are undesirable as a human protein source, they are readily consumed by other fishes 
and their role in marine ecosystems is poorly understood. It is likely that additional species will be 
discovered as a result of ongoing deepwater ichthyological surveys. 

Materials and Methods 

Measurements are straight-line, made either with a 300 mm ruler with 0.5 mm gradations (for 
total length, trunk length, and tail length), and recorded to the nearest 0.5 mm, or a 1 m ruler with 
1 mm gradations and recorded to the nearest 1 mm. All other measurements are made with dial 
calipers or dividers and recorded to the nearest 0.1 mm. Body length is head plus trunk length. 

669 



670 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 36 

Head length is measured from the snout tip to the posterodorsal margin of the gill opening; trunk 
length is taken from the end of the head to mid-anus; maximum body depth does not include the 
median fins. The jaw rictus of the paratypes were surgically cut on the right side to allow the accu- 
rate examination of dentition, a necessary procedure. Head pore terminology follows that of 
McCosker et al. (1989:257) and McCosker and Chen (2000). Vertebral counts (which include the 
hypural) were taken from radiographs. The mean vertebral formula (MVF) is expressed as the aver- 
age of predorsal, preanal, and total vertebrae. Type specimens are deposited at the California 
Academy of Sciences, San Francisco (CAS). Institutional abbreviations follow the Standard 
Symbolic Codes for Institutional Research Collections in Herpetology and Ichthyology (Leviton et 
al. 1985). 

Genus Ophichthus Ahl, 1789 

Ophiclnhiis Ahl, 1789: 5 (type species Miiraena ophis Linnaeus 1758, by original designation). 

Ophichthus piiUiis McCosker, sp. nov. 

(Figs. 1-3: Table 1) 

Material examined. — Holotype: CAS 222666, 451 mm TL, a ripe male, from Angola (12°24'S, 
13°22'E), 106-107 m, otter trawl, RA^ Dr. Fridtjof Nansen. Sta. 3608. between 1604-1634 on 1 Aug. 2005. 
Paratype: CAS 222667, 529 mm TL, a ripening female, from Angola (07°04'S, 12°00'E), 150-154 m, otter 
trawl, RN Dr. Fridtjof Nanseiu Sta. 3767, between 1615-1645, on 20 Aug. 2005. Nonparatype: unnumbered 
specimen, Centro Oceanografico de Canarias, Tenerife, 534 mm TL, a ripe male, from Bissau, precise local- 
ity and depth unknown. 

Diagnosis. — A moderately elongate species of Ophichthus with: tail 57-61%, head 
8.5-8.9%, and body depth at gill opening 2.8-3.3% of total length; dorsal-fin origin well behind 
pectoral-fin tips; pectoral fin rounded, not elongate and well-developed; posterior nostril a hole 
above the upper lip, covered by a flap that extends to or below the edge of the mouth; upper lip 
lacks barbels between anterior and posterior nostrils; pores small but conspicuous, SO 1 -i- 4, 10 4 
-I- 2. POM 6-1-2; teeth small and conical, biserial on anterior vomer and jaws; coloration uniform 
gray-brown to nearly black; total vertebrae 149-153, mean vertebral formula 20.3-56.3-151.3. 

Counts and measurements of the holotype (in mm). — Total length 451; head 38.5; 
trunk 138.5; tail 274; predorsal distance 65; pectoral-fin length 9.6; pectoral-fin base 4.05; body 
depth ca. 12.5 at gill openings; body width ca. 12.0 at gill openings; body depth at anus ca. 15; 
body width at anus ca. 14.5: snout 6.7; tip of snout to rictus 11.0; eye diameter 3.25; interorbital 
width 5.4; gill opening height 5.25; isthmus width 6.6. Vertebral formula 18-55-149. 

Description. — Body moderately elongate, subcircular to level of anus, then becoming more 
compressed, its depth at gill openings 30-36 in TL. Branchial basket moderately expanded. Head 
3.6-3.8 in trunk. Head and trunk 2.3-2.6 and head 11-12 in TL. Snout rounded, moderately acute 
when viewed from above; a short groove bisecting underside of snout nearly to tip of upper jaw; 
snout, lips and chin densely covered with minute sensory papillae. Lower jaw included, its tip 
reaching beyond anterior base of anterior nostril tube. Upper jaw not elongated, rictus behind a ver- 
tical from posterior margin of eye. Eye not enlarged, 3.2-3.6 in upper jaw and 10.1-11.9 in head. 
Anterior nostrils tubular, extending ventrolaterally from snout at ca. 20°, reaching below upper lip 
but not reaching tip of chin when directed forward. Posterior nostril a hole above upper lip, cov- 
ered by a flap that extends below the edge of mouth. There are no barbels along upper lip between 
the anterior and posterior nostrils. Dorsal-tin origin begins well behind pectoral fin about a head 
lensth into trunk lenath. Median tins low but obvious, ending in a shallow groove a little more than 



MCCOSKER: NEW DEEPWATER SNAKE EEL 



671 




FiGLTtE 1. Holotype of Ophichthus pulliis sp. nov., CAS 222666, male, 451mm. 




\ 



Figure 2. Head of holotype of Ophichthus puUus sp. nov., CAS 222666, 
male, 451mm. 



2 eye diameters before the bluntly 
pointed tail tip. Pectoral fins 
rounded, not elongate and lanceo- 
late. 

Head pores identical in posi- 
tion and number for all speci- 
mens, small but apparent (Fig. 2). 
Single median interorbital and 
temporal pores. Supraorbital 
pores 1 + 4, infraorbital pores 4 -i- 

2. lower jaw pores 6, preopercular pores 2, supratemporal pores 3. Faint rows 
of sensory papillae are visible along the nape and beneath and behind the 
mandible. Lateral-line pores apparent; 8 before gill opening in a high arching 
sequence. 54-55 before anus. 143-145 total, the last ca. the distance of the 
snout from the tail tip. 

Teeth (Fig. 3) small, conical, slightly retrorse. One central and 2 on each 
side at tip of snout, followed by an intermaxillary rosette of about 5 irregular 
pairs of teeth, followed by a single row of 8-9 small vomerine teeth, decreas- 
ing in size posteriorly. Maxillary with about 6-9 pairs of subequal irregularly 
biserial teeth, followed by 3-5 uniserial teeth. Lower jaw with about 4-6 pairs 
of irregularly subequal biserial teeth, followed by 13-18 uniserial teeth. 

Color in ethanol uniform gray-brown to black. An irregular pattern of fine 
black specks equal in size to lateral-line pores along dorsal and ventral surface 
of trunk and tail. Inner margins of lips pale, with a fine black line extending 
from beyond eye to rictus. Median fins basally pale. Anterior nostrils, tail tip, 
anal opening, lateral line and cephalic pores, and margin of median fins pale. 
Posterior pectoral fm margin pale. Peritoneum pale. Inside of mouth pale, 
densely speckled with dark brown flecks. 

Size. — The largest known specimen was 585 mm TL. It was trawled from off Angola, frozen 
and thawed and in very poor condition and ultimately discarded. 

Etymology. — From the Latin pullus, dark-colored, in reference to its appearance. 

Distribution. — Known from the type series, from 106-154 m depth, collected from off 
Angola and Guinea-Bissau. 

Remarks. — The new species appears to be very closely related to the deepwater snake eels 




Figure 3. Denti- 
tion of holotype of 
Ophichthus pullus sp. 
nov., CAS 222666, 
male, 451 mm. 



672 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 36 





0. 


pullus 


0. aphotistos 


0. cruentifer 




Mean 


Range 


Range 


Range 


TL (mm) 


— 


451-529 


480-628 


249^28 


HL/TL 


88 


85-89 


77-81 


69-87 


Head and trunk/TL 


415 


392-427 


396^06 


380^30 


Tail/TL 


585 


573-608 


594-604 


570-620 


Depth/TL 


30 


28-33 


25-29 


23-30 


DFO/TL 


175 


144-191 


123-148 


95-160 


PF length/HL 


237 


226-249 


208-326 


238-3061 


Upper jaw/HL 


317 


286-343 


311-324 


290-370 


Snout/HL 


183 


17^188 


204-216 


200-230 


Eye/HL 


92 


84-99 


94-106 


56-92 


Predorsal vertebrae 


20 


18-23 


16-19 


14-192 


Preanal vertebrae 


56 


55-58 


58-60 


56-613 


Total vertebrae 


151 


149-153 


158-162 


14^1554 



' These data reflect the removal of an irregular specimen that was previously included 
in McCosker and Chen (2000: table 1): - n=33: 3 n=31; ^ n=48 



Ophichthus aphotistos Table 1. Counts and proportions (in thousandths) of the holotype, paratype, 

McCosker and Chen ^"d nonparatype of Ophichthus pulhis as compared to three specimens of O. 

(2000) from Taiwan aplwtistos (from McCosker and Chen 2000) and 10 specimens of O. cruentifer 

and O cruentifer (fro^McCoskeret al. 1989). (TL = total length, HL = head length.) 

(Goode and Bean 
1896) of the eastern 
Atlantic. They are 
nearly identical in 
body shape and pro- 
portions, physiogno- 
my, pore conditions, 
and dentition, and are 
very similar in their 
dorsal-fin origin, pec- 
toral-fin condition, and 
coloration. Ophichthus 
pullus differs from O. 
aphotistos and O. cru- 
entifer in the length of 
its lower jaw; when 
closed, the lower jaw 
of the new species 
extends beyond the 
bases of the anterior 

nostril tubes, whereas that of the other species either falls short or does not exceed the bases of 
those tubes. The dentition of the new species is similar in appearance, location, and number to that 
of O. cruentifer (see McCosker et al. 1989: fig. 395), but differs from that of O. aphotistos (see 
McCosker and Chen 2000: fig. 3). Whereas the jaw teeth of O. pullus are biserial anteriorly and 
uniserial posteriorly, those of most O. aphotistos are biserial throughout. This however may be 
related to the size of the specimen in that the smallest paratype (480 mm TL) of O. aphotistos also 
has uniserial maxillary dentition posteriorly (McCosker and Chen 2000: 354-355). Ophichthus 
pullus further differs from O. aphotistos (Table 1) in having a shorter head, slightly more posteri- 
or dorsal-fin origin, a shorter and blunter snout, a shorter pectoral fin (paddle-shaped rather than 
lanceolate), and fewer vertebrae. Ophichthus pullus further differs from O. cruentifer (Table 1) in 
having a longer head, a slightly more posterior dorsal-fm origin, a generally smaller eye, a shorter 
and blunter snout, a shorter and broader pectoral fin. and a darker coloration (uniform tan to gray- 
ish brown, rather than dark gray to black). Ophichthus pullus attains a larger size than does O. cru- 
entifer; the known specimens of O. pullus range from 451-585 mm TL, whereas the largest of 84 
specimens of O. cruentifer examined by Wenner (1976) was 423 mm TL and the largest of 80 spec- 
imens examined by McCosker et al. (1989) was 467 mm TL. Ophichthus cruentifer occupies 
depths similar to that occupied by the new species, and O. aphotistos has been captured by trawl 
over sand and muddy substrates between 36-1350 m (McCosker et al. 1989). and was observed 
from submersibles by Wenner (1976) to be most abundant between 250-350 m, with only their 
heads exposed over sandy substrates or resting on the sediment with their bodies in S-shaped 
curves. 

The new species was also compared to specimens and descriptions of other elongate Indo- 
Pacific species of Ophichthus. The deepwater (235^90 m) western Indian Ocean O. serpentinus 



MCCOSKER: NEW DEEPWATER SNAKE EEL 673 



Seale U917) is similar in elongation and appearance, but has more vertebrae ( 163-167) and unis- 
erial mandibular teeth. Ophichthus pullus is also similar in appearance to Ophichthus exourus 
McCosker (1999), a deepwater (450-520 m) species from New Caledonia and Fiji, which differs 
in having uniserial mandibular teeth and more vertebrae (176-177). Ophichthus brachynotopterus 
Karrer (1982), known from three deepwater (355-428 m) specimens from NE Madagascar, has 
similar but more irregular biserial dentition, a much larger eye, a more posterior dorsal-fin origin 
(above the 27th-31st vertebrae), and more vertebrae (178). 

Various subgeneric lineages can be identified within Ophichthus (sensu lato), and a compre- 
hensive examination of the more than 55 valid species may result in the elevation of several sub- 
genera to generic status (McCosker 1977; McCosker et al. 1989). Based on current knowledge, the 
new species and its relatives O. aphotistos and O. cnientifer would reside in Omochelys (Fowler 
1918. originally described as a subgenus of Pisodonophis), type species Pisodonophis cruentifer 
Goode and Bean (1896). 

COMPARAXrVE MATERIAL EXAMINED. — Ophichthus apliotistos, CAS 209192, 580 mm TL (holo- 
type). and USNM 356862, 628 mm TL, and NSYSU 3657, 480 mm (paratypes). Opliiclithus bmchyno- 
toptenis. MNHN 1979-22, 442 mm TL, and 413 mm TL (MNHN 1979-23, 413 mm TL (paratypes). 
Ophichthus cruentifer, USNM 28938, 415 mm TL (lectotype), and 80 additional specimens 61-A61 mm, as 
listed in McCosker et al. (1989: 386). Opitichthus exourus, MNHN 1995-425, 520 mm (holotype), and CAS 
89552, 429 mm (paratype). Ophiclitlius serpentinus, MCZ 9200, 495 mm, holotype. 

Acknowledgments 

I wish to thank Tomio Iwamoto for collecting the type specimens; Kent Carpenter, Michel 
Lamboeuf, and Pere Oliver for inviting my participation in the FAO workshop; Eduardo Balguerias 
Guerra for assistance during the FAO workshop; Beth Herd Guy for preparing the illustrations; the 
late Eugenie Bohlke (Academy of Natural Sciences, Philadelphia [ANSP]), and the staffs of the 
California Academy of Sciences (CAS), Museum of Comparative Zoology, Harvard University 
(MCZ). Museum National d'Histoire Naturelle, Paris (MNHN), and the National Museum of 
Natural Histor}' (NMNH), Washington D.C., for advice and assistance with specimens; Alan 
Leviton for his patience and assistance; Tomio Iwamoto for reading a draft of this manuscript; and 
the volume editors, who caught a few unpardonable goofs in the typescript. 

Literature Cited 

Ahl, J.N. 1789. Dissertatio de Muraena et Ophichtho. Dissertationes Academicae Upsaliae Jiabitae sub prae- 

sidio C.P. Tliunberg 3(1): 1-12. 
Fowler, H.W. 1918. New and little known fishes from the Philippine Islands. Proceedings of t}7e Academy of 

Natural Sciences of Philadelphia 70:2-71. 
GooDE, G.B., AND T. H. Bean. 1896. Oceanic Ichthyology, a Treatise on the Deep-Sea and Pelagic Fishes of 

the World . . . with an Atlas containing 417 figures. Special Bulletin of the U.S. National History Museum 

Vol. 1. 553 pp. 
KLarrer, C. 1982. Anguilliformes du Canal de Mozambique (Pisces, Teleostei). Faune Tropicale 23:1-116. 
Leviton, A.E., R.H. Gibbs, Jr., E. Heal, and C.E. Dawson. 1985. Standards in Herpetology and Ichthyology: 

Part I. Standard symbolic codes for institutional resources collections in Herpetology and Ichthyology. 

Copeia 1985:802-832. 
McCosker, J.E. 1977. The osteology, classification, and relationships of the eel family Ophichthidae. 

Proceedings of the California Academy of Sciences ser. 4, 41(1): 1-123. 
McCosker, J.E. 1999. Pisces Anguilliformes: Deepwater snake eels (Ophichthidae) from the New Caledonia 

region, Southwest Pacific Ocean. Pages 571-588 in A. Crosnier, ed., Resultats des Campagnes 

MUSORSTOM Vol. 20. Memoires du Museum national d'Histoire naturelle 180. 



674 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 36 

McCosKER, J.E., E.B. BoHLKE, AND J.E. BoHLKE. 1989. Family Ophichthidae. Pages 254^12 in Fishes of the 
Western North Atlantic, Part 9, Vol. 1 : Orders Anguilliformes and Saccophaiyngiformes. Sears Foundation 
for Marine Research, Yale University, New Haven. Connecticut, USA. 

McCosKER, J.E., AND Y Chen. 2000. A new species of deepwater snake-eel, Ophichthus aphotistos, with com- 
ments on Neenchelys retropinna (Anguilliformes: Ophichthidae) from Taiwan. Ichthyological Research 
47(4):353-357. 

Seale, A. 1917. New species of apodal fishes. Bulletin of the Museum of Comparative Zoology, Han'ard 
College 61(4):79-94. 

Wenner, C.A. 1976. Aspects of the biology and morphology of the snake eel, Pisodonophis cruentifer 
(Pisces, Ophichthidae). Journal of the Fisheries Research Board of Canada 33(4):656-665. 



Copyright © 2005 by the California Academy of Sciences 
San Francisco, California, U.S.A. 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 

X'olume 56. No. 37. pp. 675-683. 3 figs. December 30, 2005 



Two New Species of Melastomataceae from 
Southern Mesoamerica 



Ricardo KriebeP'^, Frank Almeda^, and Armando Estrada^ 

^Department of Botany, California Academy of Sciences, 875 Howard St., San Francisco, 

California 94103-3098; e-mail rkriebel@calacademy.org; falmeda@calacademy.org. 

-Herbaria Nacional, Museo Nacional de Costa Rica, Apartado 749-1000, San Jose, Costa Rica. 

^Institiito Nacional de Biodiversidad, Apartado 22-3100, Santo Domingo de Heredia, Costa Rica 



Two new species of Melastomataceae, Blakea vemista (Blakeeae), endemic to Costa 
Rica, and Miconia dissitinervia (Miconieae), restricted to Costa Rica and Panama are 
described and illustrated. Blakea veniista is distinguished by its epiphytic, pendent 
habit, copious indument of spreading reddish-brown hairs, and paired leaves at a 
node that are commonly dimorphic in size. Miconia dissitinervia is characterized by 
a calyx that is fused in bud but ruptures at anthesis into irregular hyaline lobes. 
Distributional and phenological notes are provided together with diagnostic illustra- 
tions, photographs taken in the field, and keys to separate these species from their 
presumed closest relatives. 

Resumen 

Dos nuevas especies de Melastomataceae, Blakea venusta (Blakeeae), endemica de 
Costa Rica, y Miconia dissitinervia (Miconieae) restringida a Costa Rica y Panama 
son descritas e ilustradas. Blakea venusta muestra un caracter vegetativo poco 
comiin en el genero, sus hojas fuertemente dimorficas en tamafio por nudo; Miconia 
dissitinervia por otro lado, presenta un caracter del androceo que comparte con 
pocos congeneres, su caliz fusionado en boton y que se rompe en antesis en lobulos 
irregulares hialinos. Tambien se incluyen notas sobre distribucion y fenologia, asi 
como fotografias y claves para separar las especies de sus parientes mas cercanos. 



Two new species of berry-fruited Melastomataceae, Blakea venusta dindi Miconia dissitinervia, 
are described from Costa Rica and Panama in the Mesoamerican biodiversity hotspot (Mittermeier 
et al., 1999; Mittermeier et al. 2004). Description of a new Blakea now brings the number of 
Mesoamerican species in that genus to 34, over 75% of which are restricted to Costa Rica and 
Panama (Almeda 2000a). Addition of another species oi Miconia brings the total number of species 
in that genus for the Mesoamerican region to 163, 127 of which are also known only from Costa 
Rica and Panama. Almeda (2000a, 2000b) commented on the importance of this southern 
Mesoamerican area as a secondary center of diversity for both of these genera and predicted that 
additional taxa would come to light as remote areas were explored. Discovery of the two species 
described here suggests that continued exploration of readily accessible collecting sites throughout 
the year will continue to yield new and noteworthy taxa. 



675 



676 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 37 

Blakea venusta Kriebel, Almeda & Estrada, sp. nov. 

Figs. 1, 2D. 

Type. — Costa Rica: San Jose: Perez Zeledon, Cordillera de Talamanca, Can-etera Interamericana km 
115-116, bosque primario y secundario a la par del camino entre Division y Hortensia, 9°28'40", 83°4r25", 
1750 m, 12 Nov. 2003 (fl, fr), R. Kriebel & D. Solano 4081 (Holotype: INB!; Isotypes: CAS!, CR!, MO!). 

Frutex epiphyticus. Ramuli sicut petioli, folia subtus pedunculi setosi 1 — 3.5 mm longis indu- 
ti. Folia in quoque pari dimorpha chartacea integra apice caudato-acuminata basi obtusa vel rotun- 
data. Folia maiora: lamina 5-9.9 x 2.6-5.2 cm lanceolato-ovata, elliptica vel elliptico-ovata, 5-ner- 
vata. Folia minora: lamina 1^ x 1-2.4 cm, ovata vel suborbiculata, 3-nervata. Flores 6-meri in 
quoque nodo 1-3; pedunculi 3-12 mm longis; bracteae omnino liberae; bracteae exteriores 7-15 x 
2.5-4.5 mm, lanceolatae vel elliptico-lanceolatae; bracteae interiores 5-11 x 4-6 mm, ovato-lance- 
olatae. Calycis tubus 1-1.5 mm longus, lobis 1.5-2.5 x 3.5^.5 mm. Petala 13-17 x5-7 mm oblon- 
ga. Antherarum thecae 4x2 mm oblongae inter se cohaerentes apice minute biporosae; connec- 
tivum nee prolongatum nee appendiculatum. Ovarium 6-loculare. omnino inferum, apice papilloso. 

Epiphytic shrub with sprawling, subscandent or pendent branchlets. Young vegetative buds, 
internodes, petioles, abaxial foliar surfaces, floral peduncles and bracts densely setose with simple, 
basally barbed reddish-brown hairs, 1-3.5 mm long. Leaves at a node slightly unequal to general- 
ly very unequal in length, somewhat dimorphic to isomorphic in shape, chartaceous, sparsely vil- 
lose to glabrous adaxially, apex caudate-acuminate, base obtuse to rounded, margin entire. Large 
leaves at a node: blade 5-9.9 x 2.6-5.2 cm. lanceolate-ovate, elliptic, elliptic-ovate or elliptic- 
oblong, 5-nerved; petioles 2-7 mm long. Small leaves a node: blade 1^ x 1-2.4 cm. broadly lance- 
olate, ovate or suborbicular, 3-nerved; petioles 0.5-5 mm. Flowers spreading but not pendent, fre- 
quently hidden under subtending leaves, 1-3 flowers in each leaf axil; peduncles 3-12 mm long. 
Floral bracts foliaceous, all free from one another, typically longer than the hypanthium proper in 
length, adaxially moderately covered with spreading basally barbed hairs, margin entire; outer 
bracts 7-15 x 2.5^.5 mm. lanceolate to elliptic-lanceolate, apex acuminate, with a somewhat con- 
spicuous midvein; inner bracts 5-1 1 x 4-6 mm. ovate-lanceolate, apex acute. Calyx tube 1-1.5 mm 
long; free portions of the calyx lobes 1.5-2.5 mm long and 3.5-4.5 mm wide, broadly deltoid to 
rounded-deltoid, with each lobe terminating in a blunt reflexed callose thickening, margin entire 
and beset with gland-tipped hairs, the adaxial surface papillose and strigillose with barbellate hairs 
especially towards the apex, abaxial surface sparsely papillose and strigillose with roughened hairs 
grading into stellate hairs basally. Petals 6, 13-17 x 5-7 mm, white, oblong, obliquely rounded api- 
cally. entire and sparsely beset with gland-tipped hairs. Stamens 12; staminal filaments 6.5-8.5 mm 
long, declined to one side of the flower opposing the style, white, inconspicuosly flushed with pink 
basally; anthers 4x2 mm. yellow, laterally connate for practically their entire length, oblong and 
somewhat arcuate in dorsal view, laterally compressed, the two pores positioned 0.5-0.75 mm 
below the apex of the anther on the ventral face, connective simple. Ovary 6-locular, papillose and 
truncate apically. Style erect and somewhat incurved distally, 8.5-9.5 mm long, glabrous, distal 
half white and basal half pink; stigma punctiform. Berry globose. 8-10 x 6-8 mm, moderately to 
sparingly stellulate-furfuraceous. Seeds mostly 1 mm long, beige, narrowly pyriform. 

Distribution and Phenology. — A local species presently known only from Costa Rica 
where it occurs in cloud forests at 1300-1750 m on the Pacific slope of the Cordillera de 

Figure 1 (right). Blakea venusta Kriebel. Almeda & Estrada. A. habit; B. fully expanded flower: C. representative sta- 
mens: D. hypanthium with subtending floral bracts, (one inner bract, petals, style, and stamens removed); E. top view of 
young fruit showing stylar scar, toral ring, and calyx lobes: F. berr\-; G. representative seed: H. enlargement of primary foliar 
veins (abaxial surface): 1. enlargement of indument on cauline internodes. (A-I from Kriebel & Solano 4081.) 



KRIEBEL, ALMEDA. AND ESTRADA: NEW SPECIES OF MELASTOMATACEAE 



677 




2 mm 



S. Troyo < 



678 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 37 

Talamanca. Blakea vemista is common to abundant at both of its known locahties and grows sym- 
patrically with other rare species of Melastomataceae such as Clidemia davidsei Almeda, Blakea 
wilsoniorum Almeda, Henriettella trachyphylla Triana, Miconia cremadena Gleason and M. 
costaricensis Cogn. Collected in flower and fruit in September and November; also with fruits in 
June. 

PaRATYPES. — CosTA Rica: San Jose: Perez Zeledon. Cordillera de Talamanca, Carretera 
Interamericana, km 115-116, bosque primario y secundario a la par del camino entre Division y Hortensia, 
9°28'40", 83°41'25", 1750 m, 3 June 2003 (fr), Kriebel & Hammel 3340 (CAS, CR, INB, MO); Perez Zeledon, 
Cordillera de Talamanca, P.N. Chirripo, Estacion Santa Elena, colectado a orilla de rio y potrero, 9°23'36", 
83°35'21", 1300-1400 m, 17 Sep. 1997 (fl, fr), Alfaw 1420 (CR, INB, MO). 

Blakea venusta is readily recognized by its dense setose indument of simple, basally barbed 
reddish-brown hairs 1-3.5 mm long on young vegetative buds, intemodes, petioles, abaxial foliar 
surface, floral peduncles and bracts, leaves that are strongly dimorphic in size at each node, short 
pedunculate flowers, calyx lobes and petals marginally beset with gland-tipped hairs, and anthers 
laterally connate for practically their entire length. In the most recent key to species of Blakea in 
Mexico and Central America (Almeda, 2000a), B. venusta keys to couplet 12 next to B. 
guatemalensis and B. foliacea, clearly its presumed closest relatives on the basis of foliar dimor- 
phism, inner and outer floral bracts that are free to the base, and laterally connate anthers. The three 
species can be distingushed by the following key: 

1. Uppermost internodes, young vegetative buds, and floral peduncles densely to moderately cov- 
ered with a scurfy paleaceous indument intermixed with or sometimes replaced by ± flattened, 
roughened hairs or varying to nearly glabrous with age; leaf blades either subpeltate or bear- 
ing domatia; peduncles 1.2-5.7 cm; calyx lobe and petal margin lacking glandular hairs; con- 
nective dorsally appendaged 
2. Leaf blades subpeltate at the base, lacking inconspicuous domatia in the angles between the 
median vein and each of the two innermost veins on the abaxial surface 

B. guatemalensis Donn. Sm. 

2'. Leaf blades not subpeltate at the base, inconspicuous domatia (these often ruptured) typical- 
ly formed in the angles between the median vein and each of the two innermost veins on the 

abaxial surface B. foliacea Gleason 

r. Uppermost internodes, young vegetative buds, and floral peduncles densely covered with sim- 
ple, basally barbed reddish-brown hairs 1-3.5 mm long; leaf blades neither subpeltate nor 
bearing domatia; peduncles 0.3-1.2 cm; calyx lobe and petal margin sparsely beset with glan- 
dular hairs; connective unappendaged B. venusta Kriebel, Almeda & Estrada 

Miconia dissitiuervia Kriebel, Almeda & Estrada, sp. nov. 

Figs. 2A-C, 3. 

Type. — Costa Rica: San Jose: Turrubares, San Juan de Mata. Area no protegida. Lajas. 9°42'20"N 
84°35'13"W. elev. 600 m. 26 Nov. 1983. A. Estrada et. al. 3101 (Holotype: CR!; Isotypes: CAS!, INB!, MO!). 

Section Amblyarrhena. Frutex vel arbuscula 1-5 m altus. Ramuli obscure quadrangulati sicut 
petioli folia subtus inflorescentia hypanthiaque dense stellatis induti. Petioli 1.5-2.5 cm longi; 1am- 

FiGL'RE 2 (right). Miconia dissitinenia Kriebel. Almeda & Estrada. A. habit showing inflorescence: B. close-up of fully 
expanded flower showing reflexed petals: C. representative leaf showing abaxial surface: D. Blakea venusta Kriebel. 
Almeda & Estrada, fully expanded flower showing declinate semicircle of connate anthers. (A-C from live material of 
Kriebel et al. 5046: D from live material of Kriebel & Solano 4081.) 



KRIEBEL. ALMEDA. AND ESTRADA: NEW SPECIES OF MELASTOMATACEAE 679 




680 



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1 mm 



0.5 mm 



Figure 3. Miconia dissitinenia Kriebel, Almeda & Estrada. A. habit with infructescence; B. enlargement of stellate 
indument on foliar and hypanthial surfaces: C. berr}': D. representative seed; E. fully expanded flower; F. stamens, ventral 
view (left) and profile \iew (right): G. representative petal, adaxial surface. (A-G ixom Aguilar 4977 .) 

ina 10-35 x 5-15 cm elliptica vel elliptico-ovata. 3-5-plinervata. Panicula 8-10 cm longa multi- 
flora; flores 5-meri. pedicellis (ad amhesim) 0.25-0.5 mm longis. bracteolis 1 mm longis. Hypan- 
thium (ad torum) 2 mm longum; caly.x primum in cono apiculato clausus demum in lobos irregu- 
lares persistentes ruptus. dentibus exterioribus 0.15-0.25 mm eminentibus. Petala ca. 2 x 1 mm 
oblonga papillosa. Stamina isomorphica glabra; filamenta 1.5 mm longa; antherarum thecae 1 x 0.4 
mm angustae oblongae. poro ventraliter inclinato: connectivum nee prolongatum nee appendicula- 
tum. Stylus 3—1 mm glaber: ovarium 5-loculare et omnino apice glabro. 



KRIEBEL. .\LMEDA. AND ESTRADA: NEW SPECIES OF MELASTOMATACEAE 



Shnib to small tree 1-5 m tall; uppermost branchlets, vegetative buds, petioles, lower leaf-sur- 
faces, inflorescences, bracts, bracteoles, and hypanthia completely covered with an indument of 
stellate hairs. Leaves of a pair equal to unequal in size; petioles 1.5-2.5 cm long; leaf-blades 10-35 
X 5-15 cm. chartaceous, elliptic (sometimes broadly so) to elliptic-ovate, margin entire to incon- 
spicuously crenulate. apex acuminate to long-acuminate, base acute to long-attenuate, 3-5-plin- 
erved (excluding the ill-defined inframarginal pair) with the inner pair of primary subparallel veins 
diverging from the median vein in alternate or subalternate fashion. Inflorescence a terminal mul- 
tiflowered panicle 8-10 cm long; bracts and bracteoles linear, 1-2 x 0.25-0.5 mm, caducous; 
pedicels 0.25-0.5 imn long. Hypanthia (at anthesis) urceolate, 2 mm long to the torus; calyx tube 
0.25 mm long; calyx fused in bud but rupturing irregularly at anthesis into 2-5 persistent hyaline 
lobes; calyx teeth 0.15-0.25 mm long, narrowly triangular. Petals 5, ca. 2 x 1 mm, papillose adax- 
ially, white, oblong, rounded to emarginate apically, conspicuously reflexed at anthesis. Stamens 
10. isomorphic; filaments glabrous, complanate. 1.5 mm long; anthers 1 x 0.4 mm, yellow, linear- 
oblong, apiculate at the apex, laterally compressed and deeply ventrally channeled between the the- 
cae ventrally, 2-celled, the single pore ventrally inclined; connective simple, neither prolonged nor 
appendaged. Ovar>' 5-locular. completely inferior, globose, the apex somewhat depressed. Style 
3— I- mm long, erect, glabrous; stigma punctiform. Ben^y globose, purple at maturity, 4-5 mm in 
diameter; seeds 0.5 mm long, pyramidate, the testa muriculate to papillate. 

Distribution and Phenology. — Known only from the Pacific slope of Costa Rica, from 
Turrubares to the Peninsula de Osa south to Panama, where it has been collected at Puerto 
Armuelles on the Burica Peninsula from sea level to 600 m. Collected in flower between November 
and Januar}' and in fruit between November and June. 

PaR-\TYPES. — Costa Rica: Puntarenas: Puerto Jimenez, Agujas, 08°33'N 83°23'W, 23 Jan. 1995, 
Aguilar & Azofeifa 3710 (CAS. CR, INB, MO); Parque Nacional Corcovado, Estacion Sirena, Sendero Ollas, 
08°28'N 83°35'W, 9 Feb. 1994, Aguilar 3103 (CAS, CR, INB, MO); Parque Nacional Corcovado, Estacion 
Sirena. Sendero Espaveles. 08°28'N 83°35'W, 16 Jan. 1997, Aguilar 4977 (CR, INB, MO); west of Rincon de 
Osa. Peninsula de Osa, 9-12 Jan. 1970, Burger & Liesner 7253 (CR); along abandoned "high road" W of 
Rincon de Osa. 8°42'N 83°31'W, 4 Mar. 1985, Croat & Grayum 59849 (CAS, CR, MO); fila before Rancho 
Quemado, near Rincon, 08°42'N 83°33'W, 11 Jan. 1993, Gentry et. al. 78687 (CAS, INB, MO); cerca del rio 
Piro. Peninsula de Osa. 29 Dec. 2004, Kriebel et al. 5046 (CAS, CR, INB, MO); Aguabuena, 3 km W of 
Rincon, 800 m N of house of Henry Monge, 4 June 1993, Thomsen 361 (CR). PANAMA. Chiriqui: Burica 
Peninsula, San Bartolo Limite. 12 miles west of Puerto Armuelles, 24 Feb. 1973, Liesner 201 (CAS, CR). 

Discussion. — Miconia dissitinervia shares a number of diagnostic characters with M. centro- 
spenna of Panama. Both species have plinerved leaves, blunt calyx teeth, a completely inferior 5- 
locular ovary, unappendaged anther connectives and a punctiform stigma. They are easily separat- 
ed by the characters enumerated in the key below. Miconia dissitinenna has been misidentified in 
the past as Miconia argentea (Sw.) DC. probably because of the shared stellate indument on abax- 
ial foliar surfaces. Miconia dissitinervia differs from the latter in having plinerved vs. nerved foliai" 
venation, a punctiform stigma vs. clavate-crateriform stigma, unappendaged anthers vs. 
appendaged anthers, an irregularly rupturing hyaline calyx vs. nonrupturing regularly developed 
calyx lobes, and seeds with a muriculate or papillate testa vs. an angulate, smooth testa. 



682 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, No. 37 

Key to the Mesoamerican species of Miconia with a fused calyx that ruptures at anthesis. 

1 Flowers 4-merous 

2. Distal branches, petioles, and inflorescence densely covered with a mixture of rusty brown 
sessile-stellate and stipitate- stellate hairs; flowers with pedicels to 0.5 mm; anther pore 
somewhat ventrally inclined; connective neither prolonged nor appendaged 

M. calocoma Almeda 

2' Distal branches, petioles and inflorescence sparingly and deciduously covered with stellu- 
late hairs and/or minute glands or uppermost internodes and adaxial petiolar surface sparse- 
ly covered with smooth hairs and minute and appressed glandular hairs; flowers sessile or 
essentially so; anther pore dorsally inclined; connective appendaged dorso-basally 
3 Leaves 5-nerved; stigma not expanded: ovary (2)-3-locular 

, M. Valeriana (Standi.) Wurdack 

3' Leaves 5-plinerved; stigma capitate; ovary 4-locular M. centrodesma Naudin 

r Flowers 5(-6)-merous 

4 Abaxial leaf surface completely covered with an indumenta of stellate or stellate-lepidote 
hairs resulting in a white to reddish-white color 

5 Leaf blades 7.5-12 x 2.4-4 cm, 3-plinerved, stellate-lepidote abaxially; inflorescences 
2-2.5 cm long; petals glabrous; ovary apex densely setose around the stylar scar; seeds 
with a conspicuous spur at the wider truncate end M. centwspeiina Almeda 

5' Leaf blades 10-35 cm x 5-15 cm. 5-plinerved, stellate abaxially; inflorescences 8-10 
cm long; petals papillose adaxially; ovary apex glabrous; seeds lacking a conspicuous 

spur on the truncate distal end M. dissitinenia Kriebel. Almeda & Estrada 

4' Abaxial leaf surface variously pubescent but never completely covered with a conspicuous 
white or reddish-white indument 

6 Abaxial leaf surface minutely and deciduously glandular-puncticulate to glabrous on the 
actual surface and copiously beset with tufts of stalked-stellate hairs where the inner- 
most primary veins diverge from the median vein; ovary 3-locular 

M. mexicana (Bonpl.) Naudin 

6' Abaxial leaf surface variously pubescent but never beset with tufts of stalked-stellate 
hairs only where the innermost primary veins diverge from the median vein; ovary (4) 
5-locular 
7 Leaf blades 5-7-plinerved; flowers on pedicels 0.5-2.5 mm 

8 Leaves subsessile and clasping or sometimes with petioles l-5(-9) mm long; 
stamens dimorphic, the larger ones antepetalous . . . . M. dissitiflora Almeda 
8' Leaves with petioles 1.5-9 cm long; stamens isomorphic 

9 Young cauline internodes. petioles, and hypanthia densely covered with 
inconspicuously stalked asperous-headed hairs; inflorescence erect and 
branched basally at the node from which it is initiated; mature leaf blades 

5-plinerved M. friedmaniorum Almeda & Umana 

9' Young cauline internodes. petioles, and hypanthia densely covered with a 
lanate indument of curly or sinuate barbed or distally bifid hairs that are 
intermixed with and grade into a ground layer of shorter amorpho-pinoid 
hairs; inflorescence arcuate or pendent and branched well above the node 
from which it is initiated; mature leaf blades 7-plinerved 

M. pendula Umaila & Almeda 

1' Leaf blades 3-5-nerved; flowers sessile or essentially so 



KRIEBEL. ALMEDA. AND ESTRADA: NEW SPECIES OF MELASTOMATACEAE 683 



10 Branchlets. petioles, elevated primary leaf veins beneath, and inflorescences 
densely covered with stalked-stellate hairs; bracteoles oblong 

M. dorsiloba Gleason 

10' Branchlets, vegetative buds, inflorescences, and hypanthia sparsely and 
deciduoLisly stellulate-furfuraceous or uppermost internodes, adaxial sur- 
faces of the petioles and leaf blades sparsely covered with smooth hairs 
(gland-tipped in part) 0.5-2.5 mm long underlain with minute glandular 
hairs; bracteoles setiform or subulate-setose 

1 1 Leaf blades 6-20 x 3-8.9 cm, 5-nerved, adaxially sparsely covered with 
smooth hairs (gland-tipped in part) 0.5-2.5 mm underlain with minute 
glandular hairs; petals glabrous; stigma capitate 

M. Valeriana (Standi.) Wurdack 

11 ' Leaf blades 12-39 x 6.5-20.5 cm, 3-nerved, glabrous; petals densely 
granulose-papillose; stigma barely expanded. . M. lampwphylla Triana 

Acknowledgments 

We are grateful to Silvia Troyo (Fig. 1) and Claudia Aragon (Fig. 3) for the illustrations and 
Alan Chou and Dominique Jackson for technical assistance. The first author thanks Daniel Solano, 
Daniel Santamaria. Barry Hammel and Reinaldo Aguilar for companionship on field trips that 
resulted in some collections of the new taxa treated here. Two anonymous reviewers also offered 
most useful comments. The photographs in Figure 2 were taken by Ricardo Kriebel. 

Literature Cited 

Almeda, F. 2000a. A synopsis of the genus Blakea (Melastomataceae) in Mexico and Central America. Novon 

10:299-319. 
Almeda, F. 2000b. New Costa Rican and Panamanian Species of Miconia (Melastomataceae: Miconieae). 

Proceedings of the California Academy of Sciences, ser. 4, 52(4):33-54. 
MiTTERMEIR, R. A., N. Myers, AND CO. MiTTERMEiER. 1999. Wotspots: Earth 's Biologically Richest and Most 

Endangered Terrestrial Ecoregions. CEMEX/ Agrupacion Sieri'a Madre, Mexico City, Mexico. 432 pp. 

MiTTERMEER. R.A., RR. GiL, M. HOFFMANN, J. PiLGRIM, T BrOOKS, C.G. MITTERMEIER, J. LaMOREUX, AND 

G.A.B. FoNSECA. 2004. Hotspots Revisited. CEMEX/ Agrupacion Sierra Madre, Mexico City, Mexico. 390 
pp. 



Copyright © 2005 by the California Academy of Sciences 
San Francisco, California, U.S.A. 



684 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

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Indexes to Volume 56 

and to 
Supplements I and II 

(Compiled by Michele L. Aldrich and Alan E. Leviton). 



Index to Volume 56 for 2005 

Articles 1 through 37 and Short Communications 



Acacia greggii 88. 311 

Acanthaceae 88. 90. 607. 608. 615 

Acanthodoris 1. 16. 213-216. 218-239. 246. 251, 

255. 258. 261. 263-267: A. armata 235; A. atro- 

griseata 236: A. bninnea 213. 216. 220-221, 264; 

A. caenilescens 236; A. citrina 215, 239; A. 

columbina 227, 229; A. falklandica 213, 221-223, 

237, 239, 261, 264-265, 267; A. globosa 231, 236; 

A. hudsoni 213, 216, 223-225, 264; A. hitea 213, 

216, 225-227, 239, 264; A. metuUfera 236; A. mol- 

licella 236; A. nanaimoensis 213. 216, 227-229; A. 

nanega 213, 229-231, 236, 239, 261, 265, 267; A. 

omata 215; A. pilosa 213, 215-216, 218-219, 239, 

261; A. pilosa var pallida 215; A. pilosa var. 

albescens 215; A. pilosa var. novzealandiae 215; 

A. pilosa var. pinpurea 215; A. pina 213. 216, 

231-233, 264; A. planca 216. 237-239, 265; A. 

rhodoceras 213, 216, 233-235; A. serpentinotus 

213. 216, 235, 264; A. stellata 215; A. stohleri 231. 

233, 267; A. iichidai 236; A. vatheleti 237 
Acanthodoridinae 214 
Acanthurus monroviae 86 
Aciodoris 1-2 
Acouitia wrightii 88 
Acropya 666 
Acropyga 601-602, 604-605; A. acutiventris 604; A. 

amoldi 601-602, 604—605; A. bakwele sp. no v. 

601-602, 604-605; A. biitteli 604; A. /;o'o/w 604; 

A. paleaitica 604; A. nibescens 604; A. silvestrii 

601-602, 604-605 
Actinemys mcmnorata 91 
Actodoris 214 
Adalaria 15. 16, 213-214, 239-243, 261-265. 267; A. 

albopapillosa 241; A. ya/zHoe 213, 239-241, 

261-262. 264; A. /ove/n' 213, 239, 26-262; A. 

proxima 213, 241-243, 261-262 
A(i/or>'x hastatus (see Sargocentron) 86 
Ae^«>e5 208; A. cdbopunctata 208; A. piinctilucens 

208 
Aegiridae210, 214, 267 
Africa 28, 76, 83, 85, 115, 182-183, 275, 557, 563, 

575-576, 601, 604-605, 625, 627; Gulf of Guinea 

275, 282, 285, 291-292; See also by country; See 

also Lake Tanganyika 
Agelenidae 315 

Akidorididae 20, 214, 265, 270 
Akiodoridae 1 



Akiodorididae 1-2 

Akiodoris 1-3, 5-11, 15-16, 20, 213-214, 261-267; 

A. lutescens 1-3, 5-6, 11, 15, 20; A. salacia sp. 

nov. 1-2, 7-9, 11, 15, 20; A. salacia 213, 261-262, 

265 
Alaska 20, 53, 55, 61, 64-65; Aleutian Islands 

(Archipelago) 2, 6, 53, 55, 61, 64-65, 215; Gulf of 

Alaska 53; See also United States 
Alaskagorgia 53 
Albania 97, 129-131 
Alcyonacea 54 
Alcyoniidae 379 
Amaranthus 392-393, 394; A. deflexus 393-394; A. 

gracilis 392-393; A. hybridus 393; A. powellii 

393; A. viridis 392-394 
Amaurobiidae 305, 314-315 
Amaurobiinae 315 
Amauwbius 315-317, 326, 334; A. fenestndis 326, 

334 
Anadoridoidea 2, 16, 214 
Anapochrysa 285-286; A. africana 285-286 
Aiiciila 1 

Ancylodorididae 214 
Aucylodoris 255 
Andaman Islands 533 
Angola 669, 670-671 
Anguilliformes 669 
Ankylopteiyx 285, 287, 291-292; A. tristicta 285, 

287, 291-292; Ankylopteiyx sp. 287, 291 
Annobon 285, 290-292 
Antarctic (Antarctica) 12, 15-16, 20, 379, 382-384, 

642-643; Davis Sea 1-2, 12, 15, 21 
Anthozoa 53, 379 
Antrodiaetidae 305 
Apochrysa 285-286, 291; A. africanus 286; A. lep- 

talea 285-286, 291 
Apogon (see also Zoramia) 625-627, 631, 635; A. 

fragilis 625, 631, 635; A. gilberti 625, 631, 635; A. 

leptacanthus 625; A. peiiitiis 625 
Apogonidae 625 
aquatic ecosystems 107, 122 
Arabian Gulf 160 
Araneae 305, 337 
Archispirostreptus gigas 558 
Arctadalaria 16, 214 
Arctic Ocean 233, 245 
Arctobiinae 315 



687 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, Index 



Arctobius 315 

Annodoris 1-3, 11, 13, 15-16, 20; A. antarctica 1-2, 
11, 13, 15-16,20 

Amtzia 379 

Asia (see individual countries) 

Asterionella 119, 155 

Atalodoris 255 

Atlantic Ocean 27, 30, 53, 64, 80, 86-87, 199, 213, 
215, 240, 242, 247-248, 256, 259, 267, 379, 
380-382, 384: Drake Passage 379, 380, 381, 383; 
North Atlantic 213, 215, 240, 242, 256, 259; Scotia 
Sea 379, 382-383, 390; South Atlantic 379, 382; 
Western Atlantic Ocean 86; See Atlantic island 
groups by name (Canary, Elephant. Falkland, 
South Georgia, South Orkney, South Shetland) 

Arthrosphaera 51 A, 599 

Aulostomus strigosus 86 

Australia 23-24, 27-29, 199, 213. 229, 251. 267, 288, 
290, 295; New South Wales 24, 29, 207; South 
Australia 213, 229; Tasmania 236 

Australasian Plate 527, 530, 532-533 



B 



Bacillariophyta 129 

Baikal Drilling Programme 109 

Baltic Sea 103, 166 

Bangladesh 46, 48 

Barchatus 76 

Bataguridae31,32, 35, 37 

Bathydoris 16-18, 213, 261-267; B. clavigera 213, 

261-262, 264, 267; B. spiralis 16 
Batrachoides 76 
Batrachoididae 76 
Bayergorgia gen. nov. 380-381, 383. 386-390; B. 

vermidoina sp. nov. 380-381, 383, 386-390 
Bayesian analyses 34—35 
Beagle (HMS) 108 
Belarus 97 
Belgium 97 

Bering Sea 236, 240. 242. 256. 259 
biodiversity 50, 107, 109, 111. 117, 122. 129-130. 

138. 146, 148, 160 
Biofilms95, 162-163, 165. 167-168 
Bioko 285, 287-292; See also Sao Tome and Principe 
Blakea 675. 676. 678; B. foliacea 678; B. giiatemalen- 

sis 678; B. venusta sp. nov. 675-676. 678 
Bodiamis 28; B. oxycephalus 28 
Bomioclvysa squamosa 285, 287, 289, 291-292 
Brazil 87, 140. 158. 160. 295-296, 298-299. 301 
Brotula 80-84; B. burbonensis 82; B. clarkae 80. 82 

B. ferniginosus 83; B. flaviviridis sp. nov. 80-84 

B. jayakari 83; B. inargiiialis 83; B. nuielleri 83 



B. miilleri (see also B. muelleri) 83; B. multibarba- 
ta 80. 82-83; B. muhibartata 80-82; B. multicir- 
rata 83; B. ordwayi 80, 82; B. town-sendi 80-84 
Burma (see also Myanmar) 42, 50 



Cadlina 16, 19, 204, 213, 261-267; C. laevis 204; C. 
luteomarginata 213, 261-262, 264-265 

Calycidorididae 214 

Calycidoris 16, 213-214, 244-246, 261-265; C. 
giientheri 213, 244-246, 261, 265; C. giintheri 262 

Cambodia 34-35, 38-39, 48 

Cambrian 534, 537-538 

Canada 1-2, 7-8, 91, 124, 126, 146, 189, 220, 223, 
227, 233, 235-236, 240-242, 244, 259, 317; Bay 
of Fundy 242, 259; British Columbia 1-2, 7-8, 20, 
189, 198, 241-242, 267; Vancouver Island 220, 
223, 227, 235-236, 240, 242 

Canary Islands 669 

Caiithigaster 636 

Carabidae 275, 282 

Caribbean 86, 160 

Carlowrightia 88-90; C. arizonica 88; C. linearifolia 
88. 90; C texana 88, 90 

Caroline Islands 66-67, 72, 627, 632, 635 

Castanopsis 550; C. echidoncarpa 550; C. hystrix 550 

Cavernocymbium gen. nov. 305-306, 309-311, 
314-317, 321, 324, 326, 328-329, 331-332, 336; 
C. prentoglei sp. nov. 305, 310-311, 321, 324, 
326, 329, 332, 336; C. vetteri sp. nov. 305, 
309-311, 321, 326, 328, 331, 336 

Cavernularia 53, 61-64; C. vansyoci sp. nov. 61-63 

Central America (see Mesoamerica) 337 

Cenozoic 527. 530-535, 555-556. 644 

Ceratochiysa antica 285. 287, 291-292 

Cetonurus 27 

Chatrabus 76; C. damaranus 76 

Cheilidonichthys 28; C. ischynis 28; C. kiimii 28; C. 
spinosus 28 

Cheilodactylus 28; C. vittatus 28 

Chesterfield Islands 80-81, 83 

Chile 28, 221 

China 28, 32, 46, 48-^9, 75, 183-184, 521. 527-529, 
531, 537. 547-550. 552; Dayang River 533, 538, 
543, 547-548, 550; Dulong River 534-536, 543, 
547; Gaoligong Shan (seel also GLGS) 527-537, 
539-553, 555; Hengduan Mountains 527-534, 
539, 542, 547-548,^550-551, 554-555; Jingsha 
River 533; Lancang 539; Lancang River 533, 539, 
541, 547-548; Lhasa Terrane 531, 534-536, 538, 
543; Longchuan River 537-538, 543, 547-548, 
550-551; Longmen Mountains 531-534, 550; 



INDEX TO VOLUME 56 



Nanding River 533. 547-548; Nujiang River 
527-528. 530. 533-534. 536-537, 539-544, 
547-552. 554; Nushan Mountains 534. 539-540. 
548; Qinghai-Xizang (see also Tibetan Plateau) 
529; Sichuan 531. 539. 542; Tamai Nmai River 
539, 548; Tenasserim-Shan Block 534, 536-538; 
Tibet 531-532, 554-555; Tibetan Autonomous 
Region 535: Tibetan Plateau 527, 529-531, 539, 
547, 549-550; Xizang Zizhiqu (see also Tibet) 
535; Yarlung Zangbo (see also Brahmaputra) 
533-534, 539, 547-548, 550; Yunnan 528-529, 
533, 535, 538-539, 547, 549-550, 554-555 

Chlamydomonas 174 

Chromodorididae 16, 261 

Chrosioderma 337-338, 340-347, 350-377; C. 
albidiim 337-341, 362-363; C. analalava sp. nov. 
337, 341-342, 364; C. havia sp. nov. 337, 340, 
342, 353, 365-366: C mahavelona sp. nov. 337, 
340-341, 343. 367; C. mipentinapentina sp. nov. 
337, 340. 344, 368; C. namoroka sp. nov. 337, 
340, 345, 350, 369; C ranomafana sp. nov. 337, 
340-341, 345, 346, 352-355. 359. 361. 370-372; 
C. roaloha sp. nov. 337. 341. 345-346, 352, 356, 
360. 373: C. soalala sp. nov. 337, 340-341, 347, 
352, 357-358, 374-376 

Chnsopa (see Glenochiysa) 285 

Chnsopeiia 288, 291-292; C. congnia 288, 291-292 

Chrysopidae 285-286, 288, 291-292 

Coelotinae 305, 315-317, 336 

Coleoptera 275 

Colombia 86, 337 

Coniocompsa silvestriana 291 

Coniopterygidae 285, 291 

Conioptenx 291 

Coral Sea 83 

Corambe 16, 18 

Corambidae 16, 19-20, 214 

Coniiger spinosus 86 

Coryphaenoides 24, 27; C. annatus 27; C. denticula- 
tus 24 

Costa Rica 302, 337, 675- 676, 678, 681 

Croatia 97 

Cryogorgia gen. nov. 54-59; C. koolsae sp. nov. 
54-59 

Cryptoscenea 291 

Ctenogobiops 622 

Cuora 32, 37 

cyanobacteria 170 

Cyclemys 33, 35-37 

Cynomacrurus 27, 29; C. piriei 27 



D 



Deep Sea Drilling Project 109 

Democratic Republic of the Congo 287 

Dendrodorididae 189-190, 201-202, 204-207, 209 

Dendwdoris 189-195, 197-198, 201-204, 206-208; 
D. albobriinnea 198; D. angolensis 206; D. 
arboresceus 190, 204, 206, 208; D. areolata areo- 
Icita 206; D. areolata nigrolineata 206; D. azinae 
198; D. azineae [sic] 190; D. behrensi sp.nov. 189, 
191-192. 194-195, 197-198, 201-204, 206; D. 
coronata 195, 198, 206; D. denisoni 207; D. elon- 
gata 198, 206; D. fumata 190, 195, 199, 206, 208; 
D. gemmacea IQl; D. grandiflora 195, 206; D. 
krebsii 190, 204, 206; D. kresbsii [sic] 208; D. lim- 
bata 195, 198, 206; D. lugubris 190; D. miniata 
204, 206; D. nigra 206; D. nigromaculata 201, 
204; D. pharpa 206, 207; D. rubra var. nigromac- 
ulata 206; D. senegalensis 206; D. stohleri sp.nov. 
189-195, 198; D. tuberculosa 195; D. warta 204, 
206 

Dendronotacea 67 

Denmark 97 

Diaphorodoris 16, 19, 213-214, 246-254, 261-265, 
267; D. lirulatocauda 213, 249-251, 261-262; D. 
luteocinta 213, 246-248, 261-262, 265; D. 
luteocincta var. alba 247; D. luteocincta var. retic- 
ulata 247; D. mitsuii 213, 251-252, 261-262, 264; 
D. papillata 213, 253-254, 261-262, 264 

Diatoms 

(Excludes diatoms in checklist, pp. 400-480) 

Achnanthes 396; A. brevipes 133, 141; Achnanthes 
lanceolate 396 

Actinella penzhica 184 

Actinocyclus 133, 135, 141, 403, 495, 639, 640, 642, 
646, 648, 654; A. barronii 639, 642, 654; A. chal- 
lengeri 640; A. ingens var. nodus 640; A. jouseae 
639-640, 646, 648; A. mutabilis 642; A. nigriniae 
639-640, 646, 648; A. cf. ochotensis 135; A. prael- 
lipiticus 642; A. radionovae 639, 642-643, 654; A. 
subtilis 133; A. cf. subtilis 135 

Amphora 126, 133, 141-142, 149, 165; A. cojfeae- 
formis 133; Amphora holsatica 133, 142; A. cf. 
tomiakae 133, 141 

Anabaenopsis circularis 133 

Aneumastus 132, 142; A. albanicus 132; A. hum- 
boltianus 132; A. rosettae 132 

Anomoeoneis 396 

Aulacoseira 111, 116, 119, 124, 141; A. baicalensis 
116, 119 

Azpeitia 639, 641-643; A. biannulata 642; A. bukiyi 
641, 641, 650, 652; A. tabularis 641 



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Bacillaria 126-127, 133, 142; B. paxillifer 133 

Berkeleya scopolurum 165 

Brachysira 396 

Campylodiscus 135-136, 142; C. simulans 136; C 

thuretii var. baldjikiana 136; C. cf. simulans 

135-136, 142 
Cerataulus 133; C. tiirgidiis 133 
Ceratium 133; C. fiisiis 133: C. pulchellum 133 
Cestodiscus 640, 642-643; C. praerapax 640. 642; C. 

iimbonatiis 642 
Chaetoceros 132-133, 134, 141; C wighamii 

133-134; C. /»He//e/7 133, 141 
Cocconeis 120, 133, 142; C. placeutida 120, 133, 

142; C. scutellum 133 
Cy do Stephanos 113 
Cyclotella 113, 116, 119, 125, 132-134, 140-141, 

174; C. caspia 133; C choctawhatcJieeana 133; C 

aff. choctawhatcheeana 132-133, 134. 141; C. 

ciyptica 174; C kuetzingiana 113 
Cymbella 116, 128, 142; C stiixbergii 116, 128 
Cymboplewa 132; C. albanica 132; C. /rt/c/ var. /z//-rt 

132; C. /Hra 132 
Didymosphenia dentata 116 
Diploneis 142, 165 
Enteroniorpha 136; £. prolifera 136 
Entovjoneis 133, 142; E. paludosa 133, 142 
Epithemia 396; £. adnata var. proboscidea 396; f. 

adnata var. proboscidea 396 
£w7zo//fl 116, 119, 126, 142, 179, 183-185, 395, 397; 

£. c/f?ve/ 116, 119, 126, 179, 183-185; £. c/^ve/ 

var. baicalensis 184; £. c/^vez var. sinica 184; f. 

hispida 184; £. lacusbaikalii 183-184; £. macida- 

ta 184; £". mondon 183; £. praenipta 183; £. 

praerupta var. inflata 183 
Fallacia pygmaea 166 

Eragilaria 133, 142-144. 396; /=: cortonensis 396 
Frustidia 113, 143 
Gomphoneis herculeana 396 
Gomphonema 395, 397: G hercideaniim var. robus- 

tum 396 
Gonyaidax nionacantha 133 
Gonyiaidax 137 

Grammatophora 133, 143; G oceanico 133, 143 
Gymnodinium 133 

Gywsigma 143, 156-160, 164. 172-173; G acumina- 
tum 143, 157-158, 161; G ///rora/f 164; G o^/Z^h- 

h;» 158-159; G /wrAtr/ 158; G spenceri 157-158; 

G wormleyi 158 
Hannaea 116, 123; //. fl/c//5 116 
Hantzschia 143. 165. 172; R ivVgoto 165 
Kobayasiella micropunctata 113 
Mostogloia 133. 143, 165 



Melosira 127, 133, 141; M. nummuloides 133, 141 
Navicula 119, 120, 126, 132-134, 142-144, 148, 

165-167, 173, 395-396; M cf. besarensis 134, 

143: M oyptocephala 120, 143; M hastatula 132; 

M lacusbaicali 119; A^. parahasta 132; A^. phyllep- 

ta 144, 166-167; A^. pseudoppugnata 132; A^. .s^fl//'- 

narum 144, 166 
Naviculadicta 100 
Nitzschia 126, 132-137, 142, 144, 148, 164, 395, 397; 

A', capitellata 132, 144; M coarctata 133, 

134-135; TV. constricta 133, 135; M Jw^/a 164; A^. 

granulata 135, 144; A^. revere 133-134, 136-137, 

144; A^. 5/g/;;a 133. 144 
Oscillatoria 133 
Peridinium 133 
Pinnularia 395, 397 

Plagiotropis 164; f! angulatum 164; P. vitrea 164 
Pleurosigma 133, 144, 156-158, 160, 164, 173, 177; 

P. angulatum 133, 144, 164, 111 \ P. fonnosum 133, 

144; f: sterrenburgii 158, 160 
Pliocaenicus 116, 119; P. costatus 116, 119 
Prorocentrum 129, 133, 137; P. 7;;/ca7t5 129, 133, 137; 

P. minimum 129, 133, 137 
Psammodictyon 1 65 
Scrippsiella 133 

Staurosira 120, 133, 144; S. const mens 120, 133, 144 
Stephanodiscus 113, 116, 119, 151 
Striatella 133, 145; S. unipunctata 133, 145 
Surirella 135-136, 145; 5. biseriata 145; 5. brebis- 

sonii 145; S. fastuosa 145; 5. cf. fluminensis 135, 

136, 145; S. 7»//;//to 145; 5. oir;//^ 145; 5. cf. 

scalaris 145; S. spiralis 145; 5. splendida 145; 5. 

striatula 145 
Synedra 116, 143, 145, 396; 5. tabulata 145; 5. wif/j/- 

Zafa 145; 5. vaucheriae 116 
Tabellaria 106. 124-125, 144-145; T. fene strata 145 
Thalassionema 132-133, 145; T. nitzschioides 

132-133, 145 
Thalassiosira 480, 504, 639, 641, 643; 7: /7»A-7t/ 639, 

641-643; 7: oestrupii 641 
Trachyneis aspera 145 
Tiyblionella 165 

End Diatoms 

Dicliptera resupinata 90 

Diplopoda 557, 559, 561 

direct development 199, 201-202, 204-205, 207-209 

Discothyrea 657, 659-661, 666; D. berlita sp. nov. 

657,659-661 
DNA31-32, 34, 37. 41. 176 
Doridacea 1-2 

Doridunculus 1, 3, 11, 15-16, 20, 214 
Doriopsilla 189-190, 201-202, 204-209; D. r//Z7o/> 



INDEX TO VOLUME 56 



691 



iinctata 190, 206, 208; D. areolata 190, 204-206; 

D. areolata areolata 205; D. gemela 190, 201, 

205-207. 209: D. janaina 190, 204, 206; D. nigro- 

maculata 190. 206; D. spaldingi 190, 201-202, 

204, 206 
Doris 189, 215, 239, 241, 244, 247, 255-256, 

258-259; D. aspera 258; D. beaumonti 247; D. 

bifida 215; D. bilamellata 255; D. diaphana 258; 

D. elfortiania 255; D. flemingi 215; D. fiisca 215, 

255; D. liturata 256; D. loveni 239; D. luteocincta 

247; D. muricata 258; D. nigricans 215; D. pa///- 

(ia 255, 259; D. pilosa 215; D. pilosa stellata 215; 

D. proxima 241; D. quadrangulata 215; D. 

rocinella 215; Z). sibirica 244; D. similis 215; D. 

sparsa 215, 239; D. stellata 215; D. sublaevis 215; 

D. subquadrata 215 
Draconarius 305, 315-316, 335; D. aspinatus 315, 

335 
Drosophila 553 
Dyschoriste decumbens 90 



E 



Easter Island 288 

Echinocorambe 1, 3, 15-16, 20-21 
ecosystems 107, 122, 129, 131-132, 138, 171 
Elytraria imbricata 90 
Elephant Island 381, 382 
Eocene-Oligocene 532 
Epinephelus marginatus 86 
Equatorial Africa 289 
Ethiopia 289 
Euglena 164 
euglenoids 170 
Eurasia 530-531, 549, 551 
Eurasian Plate 527, 530-531, 533-534, 548, 552 
Europe 215, 240, 397-399, 502-503, 557; See also by 
country 



Galapagos Islands 53, 80, 190 

Galidia 572; G. elegans 572 

Gambia 289 

Germany 21, 40, 50, 84, 97, 110, 158, 170 

Gilbertiodendron 602 

Gilibelemnon 379 

Glenochrysa 285, 288, 291; G conradina 285, 288, 

291 
GLGS (see also China: Gaoligong Shan) 527-554 
Gobiidae 619 
Gondwana531,534, 552 
Goniodorididae 1, 16, 19, 20, 214 
Goniodoris 1, 16, 19-20, 213-214, 261-267; G 

nodosa 261-262, 265 
Greenland 240, 242, 256, 258-259 
Guinea Bissau 289, 669, 671 
Gymnodorididae 214 

H 

Halipteris 64 

Halobatrachus 76 

Hawaii (Hawaiian Islands) 23-24, 27-28, 76, 80-81, 

83-84, 87; Oahu 23-24, 81 
Helicoconis 291; H. (Capoconis) bazi 291 
Hemerobiidae 285-286, 289, 291 
Hemerobius: H. nairobicus 289; H. reconditus 291 
Henrya insularis 90 
Heosemys 31-35, 37-38; H. annandalii 31-32, 35; H. 

depressa 31, 35; H. grandis 31, 35; H. leytensis 

31-35, 37-39; H. spinosa 31, 35 
Heteroconis africana 29 1 
Hieremys 32; H. annandalii 32 
Himalaya(s)(an) 531, 534, 536-538, 540, 543, 549 
Holocentrus adscensionis 86 
Hungary 97, 182 

Hymenocephalus 27, 29; H. aterrimus 27 
Hymenoptera 601, 657 



Falkland Islands 120-121, 221 

Farrella elongata 10 

Fiji 80-81, 84, 288, 619, 623, 625, 627, 636, 673 

Finland 183-184 

Flourensia cemua 88 

Formicidae 601, 657 

France 199, 240, 242, 256, 259 

French Guyana 160 



Gabon 601-602, 604 
Gadiformes 23 



Iceland 240, 242, 256, 259 

India 46, 48, 50-51, 76-77, 531, 534, 548-549; 

Brahmaputra River 533, 539, 547-548, 550 
Indian Ocean 64, 76, 160, 625, 672; See island groups 

by name 
Indian Plate 527, 530-531, 533-534, 548 
Indochina 530-534, 550 
Indo-Malaya(n) (biogeographic realm) 527, 529, 

550-552 
Indonesia 40, 46, 48, 50, 533, 625, 627, 631-632, 

635; Sumatra 533 
Indo-Pacific 28, 66, 74, 80, 82, 86, 625, 672 
Ireland 97 



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Jacobinia 607-608, 615; J. leucothamna 607-608, 

615 
Japan 28, 53, 83, 182-183, 236, 251, 289; Ryukyu 

Islands 28, 623, 631, 635; Amami O Shima Islands 

631 
Johnston Island 80-81, 83 
Justicia 90, 607-615; J. cabrerae 615; J. califomica 

90; J. candicans 90; J. chol 614; /. cobensis 608; 

J. costaricana 614; J. dendropila 608; J. edgar- 

cabrerae sp. nov. 607-608, 610-611, 613-614; /. 

leucothamna sp. nov. 607-608, 610-612, 615: J. 

longii 90; J. lundellii 608, 611; J. luztnariae sp. 

nov. 607-612; J. nevlingii 614; J. salviiflora 615; 

J. sonorae 90; J. uxpanapensis 614 

K 

Korea 28 

Kuronezumia 27; K. bubonis 27 



Lake Tanganyika 115, 127, 182-183 

Lamellidoridella mitsuii 25 1 

Lamellidorididae 214 

Lamellidoris 215, 246, 256. 259, 261; L. aspera 259; 
L. bilamellata 256; L. bilamellata praecedentis 
256; L. bilamellata van liturata 256; L. bilamella- 
ta van pacifica 256; L. hystricina 259; L. varians 
259 

"'Lamellidoris n. sp. vel. van praecedentis" 256 

Larrea tridentata 88 

lecithotrophic (development) 201, 204-205, 207-208 

Lepidohynchus [sic] 29 

Lepidorhynchus 23-25, 27-29; L. denticulatus 23-25, 
27-29 

Lophodoris 1 

Lord Howe Island 28 

Loyalty Islands 80, 83 



M 



MacClade 16, 34 

Macedonia 97, 130 

Macrobuninae 315 

Macrobunus 315 

Macrouridae 23, 29 

Macrourus 24 

Madagascar 283, 288, 292, 337-338, 340-348, 557, 
558, 561, 564, 567, 570, 572, 574, 625, 627-628, 
635, 657-658, 661, 663, 665-666, 673 

Malacocephalus 27; M. laevis 27 



Malaysia 38, 46, 48^9 

Maldive Islands 635 

Mantispidae 285, 290-291 

Marionia 66-68, 72-74; M. albotuberculata 74; M. 
babai 14; M. bathycarolinensis sp. nov. 66-68, 
72-74; M. berghi 67; M. blainvillea 74; M. chloan- 
thes 74; M. cucullata lA; M. cyanobranchiata 74; 
M. dakini 74; M. echinomuriceae 14; M. fulvicola 
74; M. gramdaris 74; M. levis 74; M. olivacea 74; 
M. pambanensis 74; M. pellucida 74; M. platycte- 
na 74; M. putulasa 74; M. rubra 74; M. tessellata 
74; M. viridescens 74 

Marioniopsis 66-61 , 72 

Marshall Islands 80. 83, 625, 627, 631-632, 635 

Mauritius 657-659, 661, 666 

Mecicobothriidae 305 

Mediterranean 53, 132, 160, 247, 254, 317, 384 

Melastomataceae 675, 678 

Mesoamerica 675; See also Costa Rica, Panama 

Mexico 88-90, 189-191, 193, 195, 197, 202, 231, 
235, 317, 400, 516, 607-608, 610-611, 614-615, 
617-678; Baja California 189-191, 195, 197, 
201-202, 225, 231, 233, 235, 607; Campeche 607, 
610-611, 615: Chiapas 607, 614; Chihuahua 
88-90; Coahuila 88-89; Durango 89; Gulf of 
California 189-190, 193, 201, 269; Nuevo Leon 
88-89; Quintana Roo 607-608, 610-611, 614; San 
Luis Potosi 88; Sonora 88-90, 231, 607, 616; 
Yucatan Peninsula (also Yucatan) 607-608, 
610-611, 614-615; Zacatecas 89 

Micromus 285, 290-292; M. africanus 291; M. 
timidus 285, 290-292 

Miconia 675, 678, 680-682; M. argentea 681; M. 
calocoma 682; M. centrodesma 682; M. centros- 
penna 681-682; M. dissitiflora 682; M. dissitin- 
ervia sp. nov. 675, 678, 680-682; M. dorsiloba 
683; M. friedmaniorum 682; M. lamprophylla 683; 
M. mexicana 682; M. pendula 682; M. Valeriana 
682, 683 

Microsymplocos 296 

Miocene 532-533, 639-643 

MrBayes 35 

Molucca Islands 625, 632 

Mongolia 183 

Mozambique 287, 289, 625, 627, 635 

Muraena ophis 670 

Muriceidae 53 

Muriceides 64 

Musola impar 285 

Myanmar (see also Burma) 42-46, 48-49, 391-392, 
529, 531, 533-534, 536-539, 542, 547-550, 552, 
555; Chin Hills 48: Indo-Burman Mountain Range 



INDEX TO VOLUME 56 



693 



48; Kachin State 42-43, 48, 537, 539, 549; 

Rakhine State 42. 48 
Myripristis jacobus 86 
Myrmeleon sp. 290-291 
Myrmeleontidae 286, 290-291 
Mytilus galloprovincialis 131, 136 

N 

Neosymplocos 295-296, 299, 302 

Nepal 534 

Netherlands 64, 87, 122-123, 125, 139, 156, 158-160 

Neuroptera 285-286, 291-292 

New Caledonia 27-28, 80-81, 83, 111, 121 

New Zealand 23, 27-28, 158, 160, 177, 181, 185, 

231,236,305 
Nezumia 27; N. propinqua 27 
Norfolk Island 28 
North America 32, 91, 119, 121, 189-190, 198-199, 

213, 215, 220, 223, 227, 233, 243, 249, 295, 317, 

393 
Norway 240, 242, 256, 259 
Notiobiella 285, 290-291 
Nudibranchia 2, 21, 66, 190, 201, 213, 265 
Nuxia verticillata 661, 666 



o 



Ocean Drilling Programme 109 

Octocorallia 53 

Odontomacninis 27, 29; O. murrayi 27, 29 

Oicode spina 255 

Okenia 213-214, 261-267; O. mediterranea 

261-262, 265 
Okiodespina 255 
Oligocene 532, 642 
oligotrophic habitats 398 
Omochelys 673 
Onchidiorus 255 
Onchidora 255 
Onchidoridae 16, 265 
Onchidorididae 1, 16, 19-21, 213-214, 216, 

239-241, 243, 246, 249, 253, 258, 260-262, 265, 

267 
Onchidoridoidea 19 
Onchidoris 16, 213-215, 239, 255-261, 263, 

264-265, 267; O. bilamellata 213, 255-258, 261; 

O. bouvieri 261; O. leachii 256; O. liturata 256; 

O. muricata 213, 239, 258-260, 264 
Onchidorus 255 
Oncidiodoris 255 
Oncodoris 255 
Ophichthidae 669, 673 



Ophichthus 669-673; O. aphotistos 669, 671-673; O. 

brachynotopterus 673; O. cruentifer 669, 

671-673; O. exourus 673; O. pullus sp. nov. 

669-673; O. serpentinus 673 
Opisthobranchia 189-190, 198, 204, 207 
Optonurus 24; O. denticulatus 24; See also Lepido- 

rhynchus denticulatus 
Ordovician 537-538 
Osmylidae 291 



Pacific Ocean 20, 53, 55, 61, 80, 82, 86, 109, 160, 
189, 190, 195, 197-199, 201-202, 204-206, 208, 
215, 240, 242, 256, 259, 267, 288, 290, 380, 394, 
625, 639, 673; Eastern Pacific 53, 80, 82; 
Equatorial Pacific Ocean 639; North Pacific 215, 
240, 242, 256, 259; northeastern Pacific Ocean 53, 
55, 61; tropical western Pacific 53; Western 
Pacific 27, 53, 64; See also Pacific Islands by 
island-group names 

Palau Islands 623, 625, 627-628, 631-632, 635 

Palearctic (zoogeographic realm) 527-529 

Paleozoic 533-535, 537, 543 

Panama 50, 53, 190, 206, 675, 681 

Pangea 531, 533 

Panyaenemys subgen. nov. 31, 38 

Papua New Guinea 25 1-252, 288, 625, 627, 63 1-632, 
635 

Parablennius 87; P. pilicornis 87 

Paracis 66, 67, 72 

Paramuriceidae 53, 64 

Paramuriceinae 53 

Paratritonia 66, 67, 73-74; P. lutea 13-1 A 

Parazanomys gen. nov. 305-307, 314-317, 320, 
322-323, 326-327, 330, 335-336; R thyasionnes 
sp. nov. 305-307, 320, 322-323, 326-327, 330; P 
thyassiones [sic] 335-336 

PAUP 16, 34 

Permian 534, 536-537 

Peru 80 

Perulibatrachus 16-lS; P. aquilonarius sp. nov. 
76-78; P elminensis 16-1%; P kilbumi 76-78; P 
rossignoli 76-78 

Phanerobranchia 214, 261 

phanerobranch dorids 1, 11, 16-19, 21 

Philippine-Pacific Plates 530 

Philippine Islands 28, 31-32, 34, 37-^1, 48^9, 84, 
623, 625, 627, 635; Palawan 31, 37-39 

Phyllidiidae 189 

phytoplankton 129-130, 132-133, 136-137, 151, 
153-154, 163 

Pinus 131; f! pinea 131; P. halepensis 131 



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PROCEEDINGS OF THE CALIFORNIA AC ADEMY OF SCIENCES 

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Pisodonophis cruentifer 673 

Placobdella 9 1 ; F. omata 9 1 ; P. papillifera 91; P. par- 
asitica 91 

planktotrophic development 201, 204-205 

planktotrophic larvae 204 

Platynini 275-276 

Platynus 276, 283; P. opacipennis 276, 283 

Plexauridae 53-54, 67, 379-380, 383 

Plexaurinae 53 

Pliocene 533, 538 

poecilogony 201, 205 

Poland 93, 96-97, 122, 129, 140 

Polyceridae 214 

Polypedates 42-A3, 45^6, 48-49; P chenfui 46, 48; 
P. colleti 46, 48; P. cruciger 46, 48; P. dennysi 46; 
P. dorsoviridis 46; P. diiboisi 46; P. dugritei 46, 
48; P. eques 46, 48; P. hungfuensis 46; P. insularis 
46; P. leucomystax 46, 48; P. macrotis 46, 48; P. 
maculatus 46, 48; P. megacephalus 46, 49; P. 
mutus 46; P. naso 46; P. nigwpunctatus 46; P. 
omeimontis 46; P. otilophus 46, 49; P. pingbianen- 
sis 46; P. prasinatus 46; P. pseudocruciger 46; P. 
puerensis 46; P. yaoshanensis 46; P. zed 46; P. 
zhaojuensis 46 

Porostomata 189, 190 

Porphyridium cruentum \1A 

Precambrian 534, 536-538, 540 

Primnoidae 379 

Pristiapogon 625 

Pristomyrentx 661, 666; P. bispinosus 661 

Proceratium 657-658, 661-666; P. avioide 657, 
661-663; P avium 657-658, 661-663; P diplopyx 
663; P. google sp. nov. 657, 663-665 

Proctaporia 255; P. fusca 255 

Prodoridunculus 1-3, 15-16, 20, 214; f! gaussianus 2 

Prosopis veliitina 88 

Protoptilum 64 

Pruniis fasciculate 311 

Psectra jeanneli 291 

Pseudodorididae 214 

Pseudoclimaciella thomensis 290-291 

Pseudomallad [sic] sjostedti 292 

Pseudomallada 285, 287, 289. 291-292; f: boninen- 
sis 285, 287, 289, 291-292; /^ luaboensis 289, 
291; P. 5/o5reJn 289, 291 

Pseudonitzschia 129, 132, 134, 136-137, 144; P 5en- 
am 129. 132, 134. 136-137, 144 

Psidium cattleianum 658 

Pterapogon 625 

Ptilosarcus 64 



Q 

Quercus chrysolepis 310, 311 

R 

Rapa Island 288 

Raphia 602 

Rhacophorus 42^9, 52; P. achantharrhena 46; P. 
angulirostris 46, 49; P. annamensis 46, 49; P. 
appendiculatus 42, 46, 49; P. baliogaster 46; P. 
baluensis 46; P. barisani 46; P. bimaculatus 46, 
49; P. bipunctatus 42, 46^9, 52; P. bisacculus 42, 
46; P. calcadensis 47; P. calcaneus 46, 49; P. 
catamitus 46; P. cyanopunctatus 46; P. dennysi 
42, 46, 48-49; P. dulitensis Al , 49; P. exechopy- 
gus 46; P. /eae 42, 46, 49; P. gaM«/ 49; P. /joan- 
gliensis 47; P. htunwini sp. nov. 43-48, 52; P. /ar- 
e-ra//^ 47; P. malabaricus Al; P. margaritifer 46; 
P. maximus 42, 47, 49; P. microtympanuin 49; P. 
modestus 46; P. namdaphaensis 46; P. 
nigropalmatus Al , 49; P. notator 46; P. orlovi 46; 
P. pardalis 46, 49; P. poecilonotus 46; P. promi- 
nanus Al; P. pseudomalabaricus Al; R. rein- 
wardtii Al, Al, 49; P. robinsonii 46; P. rufipes 49; 
P. taronensis 42, 46, 49; P. translineatus 46; P. 
tuberculatus 46; P. turpes Al, 41, 49; P. variabilis 
46; P. verrucopus 46; P. verrucosus Al, 46, 49 

P/jM5: P. microphylla 88, 90; P. trilobata 311 

RNA 175, 176 

Rosgorgia 379 

Pz<£'//za 88-90; P. nudiflora 90; P. parry/ 88-90 

Russia 1^, 6, 97, 124, 183-184; Baikal 107, 109, 
111, 115-120, 182, 187; Commander Islands 1-2, 
4, 6; Kamchatka Peninsula 4, 6, 184, 186; Kuril 
Islands 2^, 6; Lake Baikal 107, 109-111, 
114-120, 122, 179, 181-185; Okhotsk Sea 1-2, 4, 
6; Siberia 110, 116, 119, 181 



Sacalia 35 

Samoa288, 625, 627, 631,635 

Sargocentron hastatum 86-87 

Sao Tome and Principe 275, 277; Principe 285-292; 

Sao Tome 115-111, 281-283, 285-292; See also 

Bioko 
Sarotheca 615 
Scincella victoriana 391 
Semidalis sp. 291 
Senegal 288-289 
Seychelles 625, 627, 635 
Siebenrockiella 31-34, 37-39; S. crassicollis 31-35, 



INDEX TO VOLUME 56 



695 



37-39; S. leytensis 37-38; See also Heosemys 

leytensis 
Silurian 537-538 
Solomon Islands 627, 632, 635 
South Africa 28, 76, 83, 213, 237, 251; Natal 289 
South America 213, 286-287, 289, 393-394; See also 

Brazil, Chile, Colombia, Peru 
Southeast Asia 530, 547, 550, 601; See also India, 

Indonesia, Malaysia. Myanmar (Burma), Nepal, 

Philippines, Thailand 
South Georgia Island 382 
South Orkney Islands 382 
South Shetland Islands 15 
Spain 97, 140,215,254,379 
Sparassidae 337 
Sphaeralcyon 379 
Sphaeromimus 557-561, 563-565, 567-574, 

578-579, 581-593, 595-597, 599; S. inexpectatus 

sp. nov. 557, 561, 563, 565, 567-568, 570, 

572-573, 593-597; 5. musicus 557-558, 561-562, 

564-565, 567-568, 570, 573, 578-579, 581-585; 

5. splendidus sp. nov. 561, 563, 565, 567-570, 

573, 586-592 
Sphaeropoeus 561, 564; 5. musicus 561, 564 
Sphaerotheriida 557-560, 563, 568, 573-574, 599 
Sphaerotheriidae 557, 573, 576 
Sphaerotherium 564, 566 
Stachyptilidae 379, 385 
Steindachneria 29-30; S. argentea 29 
Steindachneriidae 29 
Stenandrium barbatum 90 
Stenogorgiinae 53 
Straneoa 275-282; S. collatata 275-282; 5. selig- 

mani sp. nov. 275, 277-282 
Subergorgiidae 379, 384 
Sunda Shelf 37 
Sweden 97, 183 
Swiftia 64 
Symplocos 295-302; S. angulata 296, 298-299; S. 

candelabra sp. nov. 295-299; 5. glaberrima 296; 

S. microstyla sp. nov. 295, 299-302; S. organen- 

sis 301-302 



Taiwan49, 83, 669, 671 

Talinum angustissimum 88 

Tamgrinia 316 

Tanganyika Lake (see Lake Tanganyika) 

Teleostei 23, 76, 80, 619 

Tertiary 125, 182-184, 531-532, 534, 538, 555 

Tethys 548 

Tetracyclus 116, 119, 183 



Tetramerium nen'osum 90 
Thailand 48, 550 
Tharbacus 76 
Tibet (see China) 
Tonga 80, 83 

Trachemys scripta elegans 91 
Trachonurus 27 
Trachonurus sentipellis 27 
Triassic 536-538 
Tritoniidae 67 

u 

Ukraine 97 

United Kingdom 97, 169, 215; Scotland 256, 258; 
Wales 215-216, 218 

United States 54-55, 61, 88-89, 91, 146, 215, 309, 
395, 397-398; Alaska (see also Alaska [main 
heading] 198-199, 215, 227, 233, 236, 240-242, 
244-246, 249, 251, 256, 258-259; Arizona 88-91; 
California 91, 189-190, 195, 197-198, 202, 215, 
220, 223, 225, 227, 233, 240, 242, 249, 251, 256, 
259, 305, 307, 309-311, 313, 392; Colorado 317; 
Connecticut 242, 256, 259; Florida 199, 206-207; 
Great Lakes 78, 91, 119, 146-149, 151; Maine 
215, 242-243, 259-260; Massachusetts 210, 240, 
256, 258-259, 270; New Hampshire 242; New 
Mexico 88-90; Oregon 91, 199, 223, 227, 256; 
Texas 88-89; Washington 215, 220, 259, 268, 379 

Urbaniocharis 296 



Vanderhorstia 619, 621-622; V. bella 619, 621-622; 

V. mertensi 619, 622 
Vanuatu 80, 83 

Ventrifossa 29; V. atherodon 29 
Veretillidae 53, 61 
Vietnam 41, 46, 48^9 
Villiersia 255, 261, 269; V. scutigera 261 
Villiersiidae 214 

w 

Wadden Sea 157, 170-171, 174-176 
wetlands 95, 107, 129-133, 137, 141, 160 



Yap Island 627, 635 



Zanomys 305-306, 314-317, 325-326, 333; Z cali- 
fomicus 325-326, 333 



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Zargus 276; Z collatatus 276 
Zenopsis 28; Z. nebulosus 28 
Zimbabwe 287 

Zoosphaerium 558, 561, 563, 567, 569, 570, 
572-574, 599; Z hippocastanum 563 



Zoramia (see also Apogon) 625-636; Z. flebila sp. 
nov. 625-631, 635; Zfragilis 625-628, 631, 635; 
Z gilberti 625-628, 631, 635; Z leptacantha 
627-628, 631, 635; Z per//ra 627-628, 631, 635; 
Z viridiventer 625-628, 631, 633- 636 



INDEX TO VOLUME 56. SUPPLEMENT I 



697 



Index to Volume 56, Supplement I 

Biodiversity: Past Present, and Future, and The Future of Taxonomy 
edited by Nina G. Jablonski and Michael T. GhiseHn 



agriculture 106, 150-151 

ALL Species Foundation 10, 114-115, 117 

AntWeb 92, 93 

asexual species 166 

B 

bacterial communities 67-68 

bacterial genome 63, 66 

bacterial species 62-70; number of 66-67 

binomial 159-160, 176, 182, 184-185, 187, 192-193; 
maintenance of 184, 185 

biodiversity 119, 124-125, 130-132, 135-147, 149, 
15-155; biodiversity crisis 115; biodiversity loss 
87, 94, 112, 125, 135; biodiversity metrics 50; bio- 
diversity partitioning 32, 35; preserving biodiver- 
sity 130-131, 153 

biogeographic provinces 35 

biogeographic units 98, 102-103, 108 

biogeography 22-23, 68, 98, 104, 106, 111-113 

biological hotspots 87 

biological species definition 62-63, 65; See also 
species: biological species; See also species con- 
cepts 

biomes 32, 36-41, 44, 101; Seasonally Dry Biome 
37-39; tropical moist forests 151; Wetland Biome 
37^0, 42; xeric biome 39 

Brazil 154 



Cambodia 98, 101, 110, 112 

Cambrian 16, 24-31; Cambrian radiation 24, 26-29 

Carboniferous 32-39, 42, 44-49; plant biogeographic 
provinces 35 

categories 161, 163, 166, 183, 185, 197, 202, 206, 
208-212; see also species, taxonomic categories 

China 15, 25-26, 38, 98, 101-103, 107-108, 
110-112, 138, 150, 152, 154 

cladistic (Hennigianj species concept 202 

classification(s) (see also Linnaean system, Linnaean 
taxonomy, rank-free classification) 159-168, 170, 
174, 179-180, 185-188, 224; biological classifica- 
tion 162, 164, 223 

classification systems 160, 167; scientific classifica- 



tion 162 

climatic fluctuations 54, 56 

Code(s) (of nomenclature) 160, 173-175, 178, 182, 
184, 186, 193, 216, 222, 225-231; International 
Code of Botanical Nomenclature (ICBN) 216, 
226; International Code of Nomenclature of 
Bacteria 226, 228; Internafional Code of 
Zoological Nomenclature (ICZN) 160, 196, 216, 
226-227; Strickland Code 184, 216, 224 

codon bias 64 

common ancestor 159, 167 

community structure 50, 56-57, 70 

competition 12, 16-21, 41, 43 

concept of genus and species 183; see also under 
species 

conservation planning 87-88, 94, 97 

consumption 131, 150-151, 153-154 

cultural microevolution 142 



D 



Darwin, Charies 159, 162-164, 168, 200, 202, 
210-211,216,224 

data management 87, 89, 91 

database(s) 93, 116, 130, 132, 136-141, 176, 192, 
210, 227-230; biodiversity database 130, 132, 
136-137, 140; specimen databases 93; taxonomic 
databases 93, 192 

diagnosability 196-199, 207, 209 

digital imaging technology 92-93 

diversity 5, 6, 12-21, 25-27, 32, 34, 37, 40, 42, 50, 
67-68, 72-73, 75-77, 79, 81, 83, 86-89, 95, 
98-99, 101, 104, 106-109, 112; biological 13, 
98-99; dominance-diversity patterns 32, 40; pat- 
terns of 13-14, 109; theory of 12-13, 15, 16, 17, 
19,21,23 

DNA 62-63, 65-69, 71; DNA bar-coding 179; DNA 
sequencing 174, 179; transfer of DNA 63 



E 



ecomorphic replacement 42 

ecosystems 14-15, 17, 32-34, 36, 42-43, 50, 88, 106, 
111; ecosystem architecture 42; ecosystem servic- 
es 135, 138-139, 150; Late Paleozoic Ecosystems 
42 

Endangered Species Act 153 



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endemism 87, 99-101, 104, 107-109, 111; rates of 

endemism 107-108 
environmental degradation 106 
Equatorial Province 35-37 
E-Type and E-Description Initiative 117 
evolutionarily significant units (ESUs) 207, 214 
exotic species 87, 119, 121-124, 126, 128 
extinction(s) 13, 20, 21, 27, 42, 55, 87, 115, 119-126, 
131-132, 135, 137-138, 149-153, 155, 167, 198; 
extinction crisis 131, 137, 147; extinction rates 
119, 120, 121, 138; extinction-promoting traits 
125; mass extinctions 119, 127; rate of species 
extinctions 135. 151, 153 



F 



fossil taxa 185, 192 
fragmentary nature of fossils 190 



gene trees 65, 204, 215 

geographic barriers 120 

geographic differentiation 133 

Germany 154 

Global Biodiversity Information FaciUty 179 

global cooling 101 

global sustainability 130, 153-154 

global warming 152-153 

groups: monophyletic 187, 192, 196, 198, 200, 

202-207, 211; paraphyletic 162, 172, 179, 187, 

203-204; polyphyletic 187, 204 



H 



habitat deterioration 87 

habitat uniformity 120, 121 

haplotypes, mtDNA 207 

hierarchy of ranks 183; see also Linnean system 

hierarchy of taxonomic categories 208-211; see also 

taxonomic rank 
homogenization 119-126, 129 
housekeeping genes 63-65 
Hox complex 28, 30 
humans influenced species richness 56 
hunting 106, 123, 151 



identification keys 92, 114, 117 
India 150, 152, 154 
India-Eurasia collision 101 

individuality 159, 162, 164-165, 167, 216, 218, 231; 
see also species concepts 



Indochina 99, 102-104, 106-107, 109-110, 112 

Indo-Malaya 101 

Indo-Malayan realm 101-102, 107-108, 111 

infectious diseases 68 

information science 159, 170-171, 173, 175, 177, 

179, 181 
invasive alien species 152 
island biogeography 151 
isolation 13, 17-20, 71 



Jaccard's similarity index 122 



landscape partitioning 33 

Laos 98-99, 101, 104, 107, 11-113 

lateral gene transfer 62-65, 70 

latitude 14-16, 26, 37, 51, 72, 101, 107; floristic 
zonation 37; latitudinal gradient 13; Northern High 
Latitudes 35, 37; Southern High Latitudes 35, 37 

legal documents 225, 227, 229 

lineages 159, 164, 190, 196, 199-209, 211, 220; con- 
cept of a lineage 200; General Lineage Concept 
201-202, 205, 213; lineage divergence 203-204; 
population-level lineages 201, 202, 203 

Linnaean nomenclature system 136; Linnaean hierar- 
chy 136; Linnaean ranks 217; Linnaean system 
136-137, 174, 180, 183, 184, 187; Linnaean tax- 
onomy 174, 179, 182, 184 

Linnaeus, Carl 159, 162, 182, 183, 184, 185, 187, 
188, 189, 191, 193, 210, 216, 217, 222, 229 

logging 106 

loss of global biodiversity 119 

M 

Madagascar 10, 86-95, 97; fragmented habitats 88; 
predicted species distribution 94—95; species rich- 
ness map 95 

Malaysia 101, 104, 112 

mammal communities 50, 53, 55-58 

Mediterranean region 150 

megafauna 56 

metazoan radiation 24-25, 27, 29, 31 

Mexico 141, 150, 154 

Miocene 101 

mobility 21 

models of speciation 200, 201, 214 

Modem or Evolutionary Synthesis 211 

molecular evolution 140 

molecular systematics 140, 147 



INDEX TO VOLUME 56. SUPPLEMENT I 



699 



monophyly 196-199, 203, 205, 207, 209; see also 

groups 
multi-taxon surveys 10, 98, 106 

N 

natural capital 130, 135 

natural kinds 161-162, 164-165, 218, 221, 223; see 
also species concepts 

Neoproterozoic 24-27, 31; Neoproterozoic-Cambrian 
24-25 

New Pangea 119-121, 123, 125, 129 

nomen nudum 184 

nomenclatural stability 217, 220 

nomenclature 159-161, 165, 171, 174, 179, 182, 184, 
186-187, 193, 216-222, 225-231; biological 
nomenclature 216-217, 221-222; phylogenetic 
nomenclature 216-219, 221-222, 224; taxon 
names 168, 216, 218-219, 221, 224 

nucleotide sequences 140 



Palearctic 98, 101 

Paleozoic (Era) 32, 33, 35, 37, 39, 41-43, 49, 120, 
127 

Pangea 119-121, 123, 125-126, 129 

paraphyletic taxa 136 

pathogens 139 

patterns of biodiversity distribution 98 

Pax-6 28 

Permian 27, 32, 34-37, 39, 42, 49 

Permo-Carboniferous 32, 35 

phenetic relationships 136, 140 

philosophy of systematics 159, 161, 167 

PhyloCode 159, 164, 179, 187, 217-218, 221-222, 
228 

phylogenetic definition of species 65; phylogenetic 
inference 217-218; phylogenetic reality 219; phy- 
logenetic relationships 175, 196, 212, 219; phylo- 
genetic system of reference 221 

phylogeny 171, 177, 185-186, 216-218, 221, 223 

phylogeographic pattern 207 

plant-herbivore coevolution 140 

Pleistocene 50-51, 53-56, 60, 113; glacial-inter- 
glacial cycles 56 

plesiomorphic taxa 185 

poUination 139 

population differentiation 130, 132, 134-135, 
141-143 

population growth 131, 150-151 

Porcupine Cave 50-57, 59 

primary productivity 14-15 



Principle of Priority 225, 227, 229 
protostome-deuterostome ancestor 27, 29 
Provincialization 41, 44 
public health 139, 141 

R 

rank-free system (see also PhyloCode) 179, 217, 221 

ranks 159-161, 163-164, 168, 179, 182-187, 193, 
196, 206, 208-212, 217, 220-221, 223, 229; inter- 
mediate ranks 186; rank-free classifications 187 

rates of origination and extinctions 192 

refugia 104, HI 

registration of names 230 

registration system 225, 230 

reproductive isolation 196-199, 208 



seasonal climate fluctuations 101 

Silurian 32 

Sixth Mass Extinction 120; see also extinction(s) 

small herbivore richness 54 

Snowball Earth hypothesis 26, 31 

Southern High Latitude Province 37 

spatial diversity 119 

specialization 12, 16, 19, 72 

speciation 18,20,21, 104, 111 

species: biological species 165, 184, 190, 197, 198, 
199, 202, 203, 206, 208, 215; chronospecies 190, 
193; cryptic species 166, 168, 213; ecological 
species concept 197, 203; evolutionary species 
190, 193, 209, 215; paleontological species 190; 
phylogenetic species 190, 193, 197, 198, 200, 202, 
203, 206, 207, 214; reproductive barriers 199, 205, 
209, 210; sister species 208, 209; species bound- 
aries 190, 192, 197; species fusion 207, 208; 
species longevity 190; species problem 197, 199, 
203, 204, 205, 206, 210, 211, 212, 223; species 
recognition 190; stratigraphic species 190; species 
assemblages 40, 43, 90; species category 159, 166, 
167, 196, 197, 204, 205, 206, 208, 209, 210, 211, 
213; species descriptions 170, 226, 231; species 
discovery 116; species diversity 12, 15, 21, 34, 68, 
73, 83, 95, 98, 104, 109; species problem 133, 147; 
species richness 12, 21, 50, 51, 53, 54, 55, 56, 57, 
61, 76, 82, 87, 88, 89, 94, 95, 98, 99, 101, 104, 119, 
122, 146; species turnover 119; species-area curve 
123; see also asexual species, individuality, natural 
kinds, species concepts, subspecies 

species concepts 159, 161, 162, 163, 165-168, 173, 
176, 188, 190, 196-212, 214-215; definition and 
"nature" of species 132; What is a species? 130, 



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133, 134, 142; see also species (biological, 
chronospecies, cryptic, ecological, evolutionary, 
paleontological, phylogenetic, stratigraphic 
species concepts) 

subspecies 123, 135, 137, 148 

Sunda Continental Shelf 101 

Sunda Islands 101 

Sweden 154 

Switzerland 154 

systematics 130-133, 135-142, 147, 159, 161, 
167-168. 170-171, 174, 176, 179, 196, 198, 211, 
224; graduate training in systematics 142; See also 
molecular systematics 



taphonomic processes 52 

taxonomic categories 166, 208-211 

taxonomic classifications 140 

taxonomic groups 159, 164, 184, 198, 199, 210 

taxonomic information 160, 175, 192, 226; accessible 
175, 225, 229; dissemination of 160; goal of 161 

taxonomic knowledge 225 

taxonomic papers 225, 226, 227, 228, 229, 230 

taxonomic products 92, 93, 94, 118 

taxonomic rank 196, 208-209, 211-212, 217 

taxonomy 4-6, 52, 69, 75, 86-89, 91-93, 95, 114, 
116-118, 130, 139-140, 157, 159-165, 167. 168, 
170-180, 182, 184, 187-188, 192-193, 196-197, 
210-212, 216, 225-231; distributed taxonomy 
170, 175; future of taxonomy 159-160, 168, 180, 



188, 196-197, 211-212, 222; practice of taxonomy 
92, 114, 116, 118; support for taxonomy 171; tax- 
onomy web sites 170, 177; unitary taxonomy 170, 
175-180 

temperature 12, 14-18, 21, 39, 41, 43, 66, 101 

terminal taxa 217, 220 

terra firma 34, 36, 40, 42, 74, 76, 82-83 

terrestrial arthropods 72, 83, 87 

Tertiary 101 

Tibetan Plateau 101, 112 

Tree of Life 136-139, 177 

turnover 42-43, 56, 72-74, 76, 82-83, 87-88, 90 

type specimens (types) 10, 12, 15-17, 34, 37, 40, 64, 
67, 73, 92-93, 108, 114, 116-117, 173, 176, 184, 
192-193 



u 



U.S. Endangered Species Act 207 

unified concept of species 196-197, 205, 209, 211 

secondary properties 196, 203, 205 

United States 121-123, 125-126, 152-154 

urbanization 121, 128 



Vietnam 10,98-113 



w 

web sites 170, 177, 192; web-based taxonomy 
192-193; validating web-based taxonomy 193 



INDEX TO VOLUME 56, SUPPLEMENT II 



701 



Index to Volume 56, Supplement II 

Atlas of Phylogenetic Data for Entelegyne Spiders (Araneae: Araneomorphae: 

Entelegynae) with Comments on their Phylogeny 

by C.E. Griswold, M.J. Ramiez, J.A. Coddington, and N.I. Platnick 



Acanthoctenus 5, 17-18, 49, 53, 58, 60, 75, 86-87, 
95-96, 198, 216-218, 240, 246, 249, 254, 257, 
309; A. cf. spinipes 86, 216-217, 240, 246, 249; A. 
spiniger 86, 257 

Aebutina 2, 5, 21, 23-24, 49, 56, 59, 62-63, 65, 75, 
87, 95-96, 157-159, 231, 243, 251, 258, 270; 
A.binotata 23, 87, 157-159, 243, 251, 258, 270 

Africa 7-8, 14, 18-19, 24-26, 33-35, 42, 44, 87-88, 
90-92, 130-131, 134, 138, 144-145, 147, 
180-182, 220, 231-232, 236, 239, 241-242, 
244-245, 248-249, 253, 255, 259-260, 271, 278, 
288, 291, 300-303; Cameroon 7, 44-45, 92, 308; 
Cote d'lvoire 45, 92, 198, 248, 254, 262, 287, 295; 
Equatorial Guinea 92 (Bioko Island 92); Kenya 18, 
87-88; Malawi 26, 88, 135-137, 230, 232, 260; 
Morocco 26, 88, 132, 134; Namibia 33, 87, 90, 
300, 306; South Africa 7-8, 18-19, 26, 33-34, 
87-88, 90-92, 130-131, 134, 138, 144-145, 147, 
180-182, 220, 231-232, 236, 239, 241-242, 
244-245, 248-249, 253, 255, 259, 260, 271, 278, 

288, 291, 300-303; Tanzania 7-8, 44, 88, 90, 93, 
147, 150-151, 222, 225-226, 233, 244, 261, 274, 

289, 303 

Agathis australis 31, 305 

Agelenidae 1-2, 5, 8, 35, 74, 85, 290, 305 

agelenids 9, 47^8, 54-55 

agelenoids 3, 74 

Akamasia 46 

Amaurobiidae 1-3, 5, 9-10, 35, 37-38, 49, 74, 85, 

284, 294, 307 
amaurobiids 9-11, 47^9, 55, 61-62, 67, 75 
Amaurobiinae 5 
Amaurobioidea 3 
Amaurobioides 21 
Amaurobius 5, 9-11, 49, 56, 63-64, 74, 85, 94, 96, 

189, 223, 254, 257; A. fenestralis 9-10, 85, 189, 

254, 257 
Ambohima 35-36, 50, 90; A. sublima 90 
Amphinecta 12 

Amphinectidae 1-2, 5, 12, 21, 74, 85, 281, 290, 307 
amphinectids 12-13, 47^9, 61, 63, 67, 74 
Anapidae 14, 72, 86, 299 
Anisacate 12, 74 
Araneidae 2, 4, 14, 86, 139 



araneids 31, 44, 50, 54-55, 65, 67, 70 

Araneoclada 2-3, 55, 71, 72 

Araneoidea 1, 3^, 14, 32, 49, 58-59, 61, 64, 72-73, 
121 

araneoids 14-15, 43, 53, 59, 62, 72-73 

Araneomorphae 1, 3, 27, 58, 60, 71 

Araneus 4, 14, 48, 50, 54, 57, 64, 86, 94-95, 121, 
249-250, 272; A. diadematus 86, 121, 249-250, 
272 

Archaea 5, 14-15, 32, 48^9, 51-53, 55-56, 58-59, 
65-67, 72-73, 86, 95-96, 121-123, 228, 235, 250, 
266, 269, 296; A. workmani 14-15, 86, 121-123, 
228, 235, 250, 266, 269, 296 

Archaeidae 1-2, 5, 14, 72-73, 86, 121, 235, 296 

archaeids 67, 73 

Argentina 1, 15-16, 27, 39, 85-89, 91-92, 112, 115, 
118, 120, 219-220, 231, 239, 245, 249, 253, 259, 
263-264, 284, 292-293, 297-298, 302, 304, 309 

Argiope argentata 86, 139 

Argyroneta 21, 67; A. aquatica 21 

Ariadna 4, 39, 56, 58, 91, 94, 95, 118, 120, 231, 239, 
253, 259, 263, 303; A. boesenbergi 39, 91, 118, 
120, 239, 253, 259, 263; A. maxima 39, 263 

Asia 19, 35, 37; China 29, 37, 89, 91, 103, 309; India 
33, 37, 90-91, 231, 253, 255; Indonesia 90, 253, 
256 (Sumatra 90, 253, 256); Malaysia 90, 
147-149, 262 (Sarawak 90, 147-149, 262); 
Myanmar 88, 300; Nepal 37, 91, 244; Southeast 
Asia 19, 37; Thailand 37-38, 90, 157, 198, 
212-213, 233 

Australia 5, 7, 12, 14, 19, 21, 24, 28, 32, 34, 37, 
39-40, 86-87, 90-91, 108-111, 142-143, 
169-170, 172-173, 185-186, 188, 219, 221, 234, 
240-241, 248-251, 254, 256, 273, 280, 290; New 
South Wales 28, 40, 169, 170, 172-173, 248, 254, 
256; Queensland 4, 7, 28, 91, 280, 290; Tasmania 
15, 17, 86, 91, 108-111, 219, 221, 234, 249, 251; 
Victoria 28; Western Australia 90, 142-143, 
240-241, 250, 273 

Austrochilidae 1-2, 4, 15, 32, 44, 71-73, 86, 234, 
288, 299 

austrochilids 15-17, 28, 47-49, 52-54, 58-60, 68, 72, 
115 

Austrochiloidea 2, 71-72, 76 

austrochiloids 49-51, 53-55, 60-61, 67-68, 71 



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Austrochilus 15-16, 72, 86, 234, 253, 299; A. forsteri 
15, 86, 299; A. melon 16, 86, 234, 253 

B 

Badumna 5, 19-21, 56, 63, 87, 94, 96, 187-188, 223, 
254, 257, 279, 306, 311; B. Candida 19-20; B. 
longinqua 19-20, 87, 187-188, 223, 254, 257, 
279, 306 

Birrana 46 



Calilena 9 

Callobius 5, 9-10, 49, 56, 61, 74, 85, 94, 96, 190, 197, 
224, 248, 282-283, 294, 307; C. bennetti 9, 85, 
190, 197, 248, 282-283, 294; C gauchama 85, 
224; C. nevadensis 85, 307; C. pictus 9, 85 

Cambridgea 41 

Campostichomma 44^5; C. manicatum 44 

Canada 86, 121, 249-250; Ontario 86, 121, 249-250 

Canary Islands 93, 215, 240, 246 

Canoe Tapetum Clade 3, 72 

Central America 86; Costa Rica 7, 18, 41, 87, 91, 
199-201, 240, 250, 301, 307; Honduras 86; 
Panama 18, 86, 92, 147, 198, 218, 254, 257 

Cicurininae 5, 21 

Conifaber 44, 92, 302; C guarani 92, 302 

Crassanapis chilensis 86, 121 

Ctenidae 1-2, 5, 17, 75, 86, 309 

ctenids 17, 48, 53-55, 64 

Cyatholipidae 14 

D 

Deinopidae 2, 4, 18, 44, 72, 87, 236, 301 
deinopids 18, 43, 49-50, 52-53, 55-58, 66-67, 69 
Deinopis 4, 18-19, 54, 59, 62, 70, 87, 94-95, 147, 
236, 242, 248, 253, 255, 260, 272, 301; D. spin- 
osus 18, 87, 147, 236, 242, 248, 253, 255, 260, 
272. 301 
Deinopoidea 50, 60-61, 72-73, 145 
Desidae 1, 2, 5, 19, 74-75, 87, 188, 290, 306 
desids 21, 47,^9, 55-56, 61, 63, 74 
Desis 5, 13, 19-21, 48, 52, 56, 58, 65-66, 74, 79, 87, 
94. 96. 180-182, 231, 239, 253. 259, 278, 291, 
306; D. fonnidabilis 19-21, 79, 87, 180-182, 231, 
239,253,259,278,291,306 
Devendra 46 

Dictyma 5, 21-23, 47, 50, 52, 54, 56, 61, 64, 87, 
94-95, 160-161, 167, 230, 236, 246. 254-256, 
276-277, 301; D. arundinacea 87. 160-161, 167, 
230, 236, 246, 254, 256. 277; D. bostoniensis 87, 
276 



Dictynidae 1-2, 5, 21, 23, 74-75, 87, 167, 276, 277, 

301 
dictynids 21-24, 47^9, 54-56, 58-59, 61-62, 66-68 
Dictyninae 5, 21 
Dictynoidea 3 
Dionycha 3, 4 

Divided Cribellum Clade 3, 73 
Dresserus 26, 88; D. subarmatus 88 
Dysderoidea 39 

E 

Ectatosticta 29-30, 54-55, 89, 103 

Emmenomma 12, 74, 284 

Entelegynae 1-3, 50, 55-56, 60, 70, 72, 76, 242 

Eresidae 1-2, 4, 24, 50, 52, 72, 88, 134-135, 300 

eresids 24-26, 47-53, 55, 57-62, 65-67, 73 

Eresoidea 2-3, 70, 72-74, 76, 300 

Eresus 4, 25-26, 50, 62, 88, 94-95, 132-135, 232, 

259; E. cf. cinnaberinus 88, 132-134, 259; E. 

cinnaberinus 25, 88, 135, 232; E. sandaliatus 

25-26, 88, 135 
Eurasia 8, 24-25, 33, 42, 46 
Europe 2; Denmark 9, 10, 26, 85, 87-88, 135, 

160-161, 167, 189, 254, 257; Finland 87; France 

46, 93, 242, 244; Greece 25, 88, 133-134, 259; 

Italy 27, 69, 88-89, 104, 106, 236, 265, 297; Spain 

27, 88, 93, 105, 201, 215, 254-255, 257; 

Switzerland 26, 88, 135,232 



F 



Fecenia 37-38, 63, 66 

Filistata 4, 27-28, 50, 54-55, 60, 69, 88, 94-95, 

104-106, 236, 255, 265, 297; R insidiatrix 27, 69, 

88, 104-106, 236, 255, 265, 297 
Filistatidae 1-2, 4, 27, 50, 58. 64, 71, 74, 88, 227, 

297, 298 
filistatids 25, 27, 47-^8, 50, 52-58, 60-62, 65-70, 

115 
Filistatinae 4, 297 
Filistatinella sp. 115 
Fused Paracribellar Clade 3, 5, 61, 74 



Gippsicola 39 

Goeldia 5, 42, 62, 91, 94, 96, 154-156, 253, 255, 275, 

304 
Gradungula 4, 28, 55-59, 63, 65-68, 72, 89, 94-95, 

116-118; G sorenseni 28, 89, 116-118 
Gradungulidae 1-2, 4, 28, 71, 89 
Griswoldia 46 
Griswoldiinae 46 



INDEX TO VOLUME 56, SUPPLEMENT II 



703 



H 



M 



Haplogynae 2-3, 55, 60, 67, 71, 310 

haplogynes 3, 25, 39, 57 

Hersilia 67 

Hickmania 4, 15-17, 28, 54-56, 59-60, 64-65, 

67-68, 72, 86, 94-95, 108-111, 219, 221, 234, 

249, 251; H. troglodytes 15-17, 86, 108-111, 219, 

221,234,249,251 
Hickmaniidae 15 
Holarchaeidae 73 
Huntia 46 
Huttonia 5, 28-29, 48^9, 51-53, 56, 58-59, 65, 67, 

72-73, 89, 95-96, 124-125, 235, 250; H. palpi- 

manoides 28-29, 89 
Huttoniidae 1-2, 5, 28-29, 73, 89, 235 
hultoniids 29, 67 
Hypochilidae 1-2, 4, 29, 89, 296 
hypochilids 16, 29-30, 47^8, 52-55, 57, 60, 62, 67, 

69 
Hypochilus 4, 29-30, 49-50, 53-59, 62, 64-65, 68, 

71, 89, 94-95, 102-103, 231, 233, 244, 251, 265, 

267, 296; H. kastoni 89, 296; H. pococki 30, 89, 

102-103, 233, 244, 251, 265, 267, 296; H. sheari 

89,231 
Hyptiotes 43, 92, 221 



Ischalea 40-41 
Israel 42 



I 



K 



Kilyana 46 

Knikt 46 

Kukulcania 4, 7, 27-28, 50, 54-55, 67, 76, 88, 94-95, 
107, 115, 219, 227, 244, 253, 255, 267-268, 
297-298; K. hibemalis 27, 67, 76, 88, 107, 115, 
227, 244, 253, 255, 267-268, 297-298 



Lathys 5, 21-23, 56, 87-88, 94, 96, 164-165, 167, 
230, 232, 248, 253, 256, 261, 276-277; L. humilis 
23, 88, 164-165, 167, 248, 253, 256, 261, 277; L. 
immaculata 88, 230, 232, 276 

Linyphiidae 14, 72 

Liocranoides 65, 91; L. unicolor 91 

Liphistius 54-55, 67 

Loxosceles 55 

Lycosoidea 3, 37-38, 41, 46-^8, 50-51, 60, 66, 75, 
312 



Macrobuninae 5, 74, 284 

Macrobunus 5, 9-11, 64, 74-75, 85, 94, 96, 195-196, 
239, 253, 259, 284, 294; M. cf. multidentatus 9-10, 
85; M. multidentatus 195-196, 239, 253, 259, 284 

Macrogradungula 28, 72; M. moonya 28 

Madagascar 7, 14-15, 35, 40, 42, 44^5, 86, 88, 90, 
93, 121-123, 198, 205, 228, 235, 242, 244, 250, 

254, 257, 266, 269, 274, 287, 295-296, 308 
Malaika 35 

Malkaridae 72-73 
Ma//o5 21-22, 88, 221 

Maniho 5, 12-13, 64, 68, 85, 94, 96, 176-177, 183, 
248, 257, 281, 290; M. ngaitahu 12, 85, 176-177, 

183, 248, 257, 281, 290; M. pumilio 85; M. tigris 
85 

Manjala 38 

Matachia 5, 19-21, 48^9, 56, 60, 68, 74, 87, 94, 96, 

184, 188, 254, 257, 306; M. australis 19, 87, 184, 
188, 254, 257; M. marplesi 87 

Matachiinae 5 
Mauritius 40 
Mecysmaucheniidae 73 
Mecysmauchenius 67 

Megadictyna 5, 32-33, 48, 50, 59-60, 62, 64, 66, 70, 
73, 75, 89, 94-95, 139-141, 223, 238, 242, 253, 

255, 262, 272, 304; M. thilenii 32, 89, 139-141, 
223, 238, 242, 253, 255, 262, 272, 304 

Megateg 46 

Menneus 4, 18-19, 50, 87, 94-95, 144-145, 236, 241, 

245, 249, 301; M. camelus 18, 87, 144-145, 236, 

301 
Mesothelae 54, 71 
Metagonia 72 
Metaltella 5, 12-13, 65, 68, 74, 85, 94, 96, 178-179, 

183, 225, 254, 256, 261, 280-281, 292-293, 307; 

M. wmlenta 12, 85; M. simoni 12, 85, 178-179, 

183, 225, 254, 256, 261, 280-281, 292-293, 307 
Metaltellinae 66, 86, 265 
Metepeira atascadero 86 
Mexico 41, 44, 86, 89, 91, 93, 126, 139, 202-203, 

229, 231, 238, 247-248, 253, 295; Baja California 

86, 139; Chiapas 91, 93, 202-203, 231, 247, 253; 

Guanajuato 86, 238, 248; Hidalgo 93, 295; 

Tampico89, 126,231 
Miagrammopes zenzesi 92, 302 
Micropholcommatidae 14, 73 
Mimetidae 1-2, 5, 30, 72-73, 89, 243 
Mimetus 5, 30-31, 48-52, 58-59, 64, 66-67, 72, 89, 

95-96, 126-127, 229, 231, 243, 250, 258, 270; M. 

hesperus 30, 89, 126-127, 229, 231, 243, 250, 258, 



704 



PROCEEDINGS OF THE CALIFORNIA AC ADEMY OF SCIENCES 

Fourth Series, Volume 56, Index 



270 
Misionella mendensis 89, 297 
Miturgidae 46 
Mygalomorphae 67, 71, 78 
Mysmenidae 14 

N 

Namaquarachne 36, 90, 147; A^. tropata 36, 90, 147 

Neocribellatae 60, 62, 68, 71 

Neocribellate28, 71 

Neolana 5, 31, 49, 62, 89, 94, 96, 168-169, 233, 

254-256, 279, 305; N. dalmasi 31, 89, 168-169, 

233, 254, 256, 279, 305 
Neolanidae 1-2, 5, 31, 74, 89, 169, 305 
neolanids47-48, 61, 67 
Neoramia 5, 9, 49, 60-61, 74, 85, 94, 96, 174-175, 

183, 254, 256, 262, 280, 290, 305; A^. sana 9, 85, 

174-175, 183, 254, 256, 262, 280, 290, 305 
Nephila 19, 91, 220 
Nesticidae 14, 72 
New Caledonia 19 
New Guinea 32, 37, 42, 91, 210-211, 213, 240, 246, 

250, 265, 268 
New Zealand 4, 7-9, 12, 18-19, 24, 27-29, 31-32, 

34, 37-38, 40, 85, 87, 89-91, 93, 116-118, 

124-125, 139-141, 168-169, 171, 174-177, 

183-184, 188, 208-209, 223, 233, 235, 238, 

242-243, 248, 250-251, 253-258, 262, 272, 

279-281,285,290,304-306 
Nicodamidae 1-3, 5, 32, 71, 73-74, 89, 139, 304, 315 
nicodamids 32-33, 47^9, 64, 71, 73, 315 
Nicodamus 5, 32-33, 48, 50, 56-57, 59, 64-65, 73, 

90, 94-95, 142-143, 240-241, 250, 273; N. 

mainae 32, 90, 142-143, 240-241, 250, 273 
Nigma 5, 21-23, 47-48, 52, 56, 88, 94-95, 162-163, 

167, 256, 277; N. linsdalei 88, 162-163, 167, 256, 

277 
North America 8, 29-30, 44-46 



o 



Octonoba 4, 43^W, 92, 94-95, 145, 248, 253; O. 

octonarius 43; O. octonaria 43, 92, 145, 248, 253 
Oecobiidae 1-2, 4, 33, 67, 72, 90, 227, 288, 300 
oecobiids 33-34, 47^8, 51-53, 55-56, 58, 62-63, 

65-68 
Oecobius 4, 33-34, 50, 53, 58-59, 66-68, 70, 72, 90, 

94-95, 128, 227, 230, 236, 241, 271; O. navus 33, 

90, 128, 236, 241, 271 
Oonops 67 
Orbiculariae 2-A, 18, 30, 32, 47^9, 58, 62, 69-70, 

12-13, It, 147,310-311 
Oval Calamistrum clade (OC clade) 2, 75-76 



Palaeocribellatae 2 

Palpimanidae 29, 73 

palpimanids 29, 73 

Palpimanoidea 2-3, 5, 14, 29, 57, 71-73, 310 

palpimanoids 47-49, 52, 62, 67, 72-73, 315 

Palpimanus 61 

Papua New Guinea 37, 91, 210-211, 213, 240, 246, 

250, 265, 268 
Paramatachia 20 
Pararchaea 5, 34-35, 49, 51-53, 56, 58-59, 64-67, 

72, 90, 95-96 
Pararchaeidae 1-2, 5, 34, 72-73, 90 
Phanotea 46 

Philoponella cf . fasciata 92, 302 
Phryganoporus 5, 19-21, 56, 63, 87, 94, 96, 185-186, 

188, 290, 311; P candidus 19-20, 87, 185-186, 

188, 290 
Physocyclus 55 
Phyxelida 5, 35-36, 49-50, 64, 90, 94, 96, 147, 

150-151, 222, 233, 244, 261, 274, 289, 303; P 

bifoveata 90, 274, 289; P tanganensis 35-36, 50, 

90, 147, 150-151, 222, 233, 244, 261, 274, 303 
Phyxelididae 1-3, 5, 35, 50, 74, 90, 147, 274, 289, 

303 
phyxelidids 9, 35-36, 47^9, 59-61, 63, 70 
Phyxelidinae 35, 303 
Phyxelidini 63, 303 
Pillara 5, 40-41, 91, 94, 96, 169, 172-173, 311; P 

griswoldi 40, 91, 169, 172-173 
Pimoidae 14 
Pimus 5, 9-11, 49, 59, 74, 85, 94, 96, 191-192, 197, 

224, 236, 248, 254, 257, 259, 262, 282-283; P 

napa 85, 197, 236, 259; P pitus 9, 85 
Poaka 2, 5, 9-10, 37-38, 49, 75, 90, 95-96, 208-209, 

243, 251, 258, 285; P graminicola 37-38, 90, 

208-209,243,251,258,285 
Pritha nana 89, 297 
Prithinae 50, 297 

Progradungula 28; P. carraiensis 28 
Psechridae 1-2, 5, 10, 37-38, 75, 90, 309 
psechrids 37-38, 53, 55, 61, 63, 66 
Psechms 5, 37-38, 50, 60, 63, 66, 90-91, 95-96, 157, 

198, 210-213, 233, 240, 244, 246, 250, 265, 268, 

309; P argentatus 91, 210-211, 213, 240, 246, 

250, 265, 268 



R 



Raecius 5, 44^6, 58, 60, 64, 67, 92-93, 95-96, 198, 
206-207, 225-226, 248, 254, 262, 287, 295, 308; 
R. asper 45, 67, 92, 308; R. congoensis 46, 67, 92; 



INDEX TO VOLUME 56, SUPPLEMENT II 



705 



RJocqiiei 45. 67. 92, 198, 206-207, 248, 254, 262, 
287, 295; R. scharffi 44, 93, 225-226 

Retiro 5, 9-n, 48-49. 58, 74. 85. 94, 96, 193-194, 
197. 254. 257. 282-283 

RTA Clade 2-3. 74-75 

Riibrius 12, 284 

Russia 87, 230, 254, 256. 277 



Sahastata 50 

Scytodes 55 

Sege stria 39 

Segestriidae 1, 2. 4. 39. 71. 91, 303 

South America 9, 12, 21, 42; Brazil 23-24; Chile 7-9, 
12. 15-16. 19, 39, 42, 85-86, 92, 113, 115, 121, 
154-156, 195-196. 234. 239, 253, 259, 263, 265, 
267, 284, 288, 294, 299; Colombia 92; Ecuador 23, 
86-87, 92, 157-159, 218, 231, 243, 251, 258, 270; 
Peru 9, 18, 85-87, 91, 193-194, 197, 216-217, 
240, 246, 249, 253-255, 257, 275, 282-283; 
Uruguay 19, 87, 279 

Stegodyphus 4, 24-26, 62, 88, 94-95, 134-138, 230, 
232, 236, 245, 253, 255, 260, 271, 300; S. dumico- 
la 88, 271. 300; S. mimosarum 25-26, 88, 
134-138, 230, 232, 236, 245, 253, 255, 260, 300 

Stenochilidae 28-29, 73 

Stiphidiidae 1-2. 5, 40^1, 74-75, 91, 169, 290, 305 

stiphidiids 40-41, 47^8, 53, 61-62, 66-67 

Stiphidioids 3, 74 

Stiphidion 5, 40-41, 53, 91, 94, 96, 169-171, 248, 
254, 256, 280, 290, 305; S. facetum 40, 91, 169, 
170-171, 248, 254, 256, 280, 290, 305 

Symphytognathidae 14 

Symphytognathoidea 14 

Synaphridae 14 

Synotaxidae 14 



Takeoa 46 

Tama 67 

Tegenaria domestica 9 

Tengella 5, 41, 44, 59, 64, 69, 75, 91, 95-96, 
199-201, 240, 250, 307; T. radiata 41, 91, 
199-201, 240, 250, 307 

Tengellidae 1-2, 5, 37, 41, 65, 75, 91, 201, 307 

Tetragnathidae 14, 72, 91 

Textricellidae 73 

Thaida 4, 15-17, 19, 28, 49-50, 53-56, 60, 62-63, 
67-70, 86, 94-95, 112-113, 115, 219-220, 
234-245, 249, 259, 264, 267, 288, 299; T. pecu- 
liaris 15-16, 86, 112-113, 115, 219-220, 234, 245, 



249, 259, 264, 267, 288, 299 

Thaididae 15 

Theridiidae 14, 72 

Theridiosomatidae 14 

Titanoeca 5, 42^3, 62, 64, 92, 94, 96, 152-153, 156, 
222, 244, 253, 255, 262, 275, 289, 304; T. albo- 
maculata 42, 82; T. americana 42, 92, 152-153, 
156, 244, 253, 255, 262, 275, 289; T. nigrella 42, 
92, 153, 156, 222, 304; T. sUvicola 43 

Titanoecidae 2, 5, 35, 42, 64, 74, 91, 156, 275, 289, 
304 

titanoecids 42, 43, 47-48, 62-63, 65 

titanoecoids 2-3, 5, 74 

Tricholathys 5, 21, 22-23, 48, 56, 88, 94-95, 
166-167, 232, 254, 256, 276; T. spiralis 88, 167, 
254, 256, 276 

Tricholathysinae 5, 21 

u 

Uduba 5, 44^6, 50, 58, 60, 93, 95-96, 198, 205, 242, 
244, 254, 257, 287, 295, 308; U. dahli 93, 295; U. 
madagascariensis 93, 287, 308 

Uliodon 46, 64, 93 

Uloboridae 2, 4, 23, 43, 72, 92, 302, 307 

uloborids 19, 43^4, 47, 49-50, 53-55, 59, 61, 65, 
67-70 

Uloborus 4, 23, 43^4, 58, 92, 94-95, 147, 220, 238, 
241, 246, 249, 253, 255, 302; U. diversus 92, 238, 
253, 255; U. glomosus 92, 241, 246, 249; U. trilin- 
eatus 92, 147 

United Kingdom 88, 164-165, 167, 248, 253, 256, 
261, 277; Scotland 87, 236, 246 

Uroctea 4, 33-34, 53, 57-59, 66-67, 90, 94-95, 
130-131, 231, 253, 255, 260, 271, 288, 300 

Uroecobius 67 

USA 1, 5, 9, 12, 18, 27, 33-34, 42-43, 85-93, 
102-103, 107, 115, 127-128, 145, 147, 152-153, 
156, 162-163, 166-167, 178-179, 183, 187-188, 
190-192, 197, 204, 221-227, 230-233, 236, 238, 
241-244, 246, 248-251, 253-262, 265, 267-268, 
270-272, 275-277, 280-283, 287, 289, 294, 296, 
301, 304, 306-309; Arizona 44, 87-89, 92-93, 
127, 153, 156, 221-222, 243, 258, 270, 304, 308; 
Arkansas 88, 92, 230, 232, 248, 253, 276; 
California 9-10, 12, 19, 27, 34, 43, 46, 85-90, 
92-93, 128, 162-163, 166-167, 178-179, 183, 
191-192, 197, 224-227, 230, 232, 236, 238, 248, 
253-254, 256-257, 259, 262, 277, 280-283, 287, 
296, 301, 307, 309; Florida 18, 27, 87-88, 107, 
115, 147, 227, 236, 242, 244, 248, 253, 255, 260, 
267, 272; Georgia 34, 88, 90, 236, 241, 268; 
Hawaii 87, 187-188, 254, 257; Lousiana 85; 



706 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series, Volume 56, Index 



Maine 85, 282-283, 294; Minnesota 87, 276; 
Missouri 92, 145, 152-153, 156, 244, 253, 255, 
262, 275, 289; Nevada 89, 250; North Carolina 30, 
88-89, 92, 102-103, 231, 241, 246, 249, 265, 296; 
South Carolina 92; Tennessee 89, 91, 233, 244, 
251, 267; Texas 93, 204, 259, 287; Washington 
167; Washington D.C 34, 90, 128, 271; West 
Virginia 85, 190, 197, 248 



Vidole 35-36, 90, 303; V. capensis 90, 303 

Vidoliini 36, 63 

Vytfutia 5, 35-36, 60, 63, 90, 94, 96, 147-149, 253, 

256, 262; V. bedel 90, 253, 256; V. pallens 36, 90, 

147-149, 262 



w 



Waitkera 44 
Wajane 25 



Xenoctenus 65, 92 



Xevioso 5, 9, 35-36, 49, 60, 62, 66, 90, 94, 96, 232, 
242, 244, 248, 271, 303; X. arnica 36, 90, 232, 242, 
244, 248, 271, 303; X. orthomeles 90, 303 



Zorocrates 5, 44-46, 64, 75, 93, 95-96, 202-204, 
231, 247, 253, 259, 287, 295, 308; Z. cf. mistus 45, 
93,202-203,231,247,253 

Zorocratidae 2-3, 5, 44, 46, 50, 75, 92-93, 287, 295, 
308 

zorocratids 44, 48, 50, 58, 60, 64, 67, 75 

Zorocratinae 44, 46 

Zowdictyna 44 

Zoropsidae 2, 5, 44, 46, 64, 75, 93, 201, 309, 313 

zoropsids 48, 53, 55, 61, 64 

Zoropsis 5, 44, 46^7, 49-50, 53, 58, 60, 64, 67, 75, 
93, 95-96, 201, 215, 226, 240, 242, 244, 246, 254, 
257, 287, 309; Z cyprogenia 46; Z media 93; Z 
nishimurai 46; Z rufipes 47, 93, 215, 240, 246; Z 
spinimana 46-41, 93, 201, 215, 226, 242, 244, 
254, 257, 287, 309 

Zosis 92 



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9MrTH90NIAN INSTFTUTION LIBRARIES 





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TABLE OF CONTENTS 

J.P. Kociolek: a Checklist and Preliminary Bibliography of the Recent, Freshwater Diatoms of 
Inland Environments of the Continental United States 395 

George Chaplin: Physical Geography of the Gaoligong Shan Area of Southwest China in 
Relation to Biodiversity 527 

Thomas Wesener and Petra Sierwald: The Giant Pill-Millipedes of Madagascar. Revision 
of the Genus Sphaeromimus, with a Review of the Morphological Terminology (Diplopoda, 
Sphaerotheriida, Sphaerotheriidae) 557 

John S. LaPolla and Brian L. Fisher: A Remarkable New Species oiAcropyga 

(Hymenoptera: Formicidae) from Gabon, with a Key to the Afrotropical Species 601 

Thomas F. Daniel, German Carnevali, and Jose L. Tapia Munoz: New and Reconsidered 
Mexican Acanthaceae XI: Justicia in the Yucatan Peninsula 607 

David W. Greenfield and Kenneth R. Longenecker: Vanderhorstia bella, a New Goby 

from Fiji (Teleostei: Gobiidae) 619 

David W. Greenfield, Ross C. Langston, and John E. Randall: Two New Cardinalfishes 
of the Indo-Pacific Fish Genus Zoramia (Apogonidae) 625 

John A. Barron: Scanning Electron Microscope Studies of Some Early Miocene Diatoms 
from the Equatorial Pacific Ocean with Descriptions of Two New Species, Actinocyclus 
jouseae Barron and Actinocyclus nigriniae Barron 639 

Brian L. Fisher: A New Species of Discothyrea Roger from Mauritius and a New Species 
of Proceratiitm Roger from Madagascar (Hymenoptera: Formicidae) 657 

John E. McCosker: A New Species of Deepwater Snake Eel, Ophichthus puUus 

(Anguilliformes: Ophichthidae), from Angola and Guinea-Bissau 669 

Ricardo Kriebel, Frank Almeda, and Armando Estrada: Two New Species of 

Melastomataceae from Southern Mesoamerica 675 

Index to Volume 56 (including Supplements) 685